J Exp Biol-1973-WEIS-FOGH-169-230 PDF

7. Exp. Biol. (1973). 59.
169-230 With 23 text-figures Printed in Great Britain
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QUICK ESTIMATES OF FLIGHT FITNESS IN HOVERING ANIMALS, INCLUDING NOVEL MECHANISMS FOR LIFT PRODUCTION
BY TORKEL WEIS-FOGH
Department of Zoology, Cambridge CBz ^EJ, England (Received 11 January 1973) INTRODUCTION
In a recent paper I have analysed the aerodynamics and energetics of hovering hummingbirds and DrosophUa and have found that, in spite of non-steady periods, the main flight performance of these types is consistent with steady-state aerodynamics (Weis-Fogh, 1972). The same may or may not apply to other flapping animals which practise hovering or slow forward flight at similar Reynolds numbers (Re), io a to io*. As discussed in that paper, there are of course non-steady flow situations at the start and stop of each half-stroke of the wings. Moreover, it does not follow that all hovering animals make use mainly of steady-state principles. It is therefore desirable to obtain as simple and as easily analytical expressions as possible which should make it feasible to estimate the forces on the wings and the work and power produced. In this way one may make use of the large number of observations on freely flying animals to be found in the scattered literature. It may then be possible to identify the deviating groups and to approach the problems in a new way. This is the main purpose of the present studies, which both include new material and provide novel solutions. Major emphasis must be placed on simplicity. This involves approximations since the true flight system is so complicated as to be unmanageable. However, when we confine ourselves to free flight and make use of the most reliable flight data available, the task is neither as difficult nor the conclusions as unrealistic as one would expect, since it is possible to introduce simple corrections. Although entitled ' Quick estimates', this does not mean that the approach is superficial, but rather that a procedure has been devised whereby the flight performance of a given animal can be evaluated quantitatively as well as qualitatively on the basis of only a few accurate observational data and a minimum of computation. In other words, given reliable information about bodily dimensions and wing-stroke parameters, the method enables one quickly to arrive at a first-order approximation so as to assess whether the animal makes use of well-established mechanisms or employs unusual or novel principles. The following procedure is, in essence, the strategy of the present investigation. From measurements of the size and shape of the wings, the geometry of the wing stroke, the frequency of the wing beat and the weight of the animal which is sustained in hovering flight one can calculate, on the basis of steady-state aerodynamics, the minimum coefficient of lift which must be ascribed to the wings. From the same measurements one can also calculate the Reynolds number under which the wings operate,
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TORKEL WEIS-FOGH
Vertical
w
Horizontal Chord, c(r)
Fig. 1. Simplified diagram of normal hovering flight. (A) The instantaneous forces. (B) The animal seen horizontally from the side, and (C) vertically from above.
and from this figure one can obtain (using published data on wings) the maximum coefficient of lift which may be expected under these conditions. If the minimum coefficient of lift for steady-state aerodynamics does not exceed the maximum coefficient of lift obtainable at the Reynolds number in question one assumes that steadystate aerodynamics are adequate to explain hovering flight, and from then one can go on to calculate the power requirement and other parameters of the flight mechanism. If, on the other hand, the minimum coefficient of lift for steady-state aerodynamics is greater than the maximum obtainable, it is clear that hoveringflightcannot be explained by steady-state aerodynamics and that a new approach must be made.
AERODYNAMIC RELATIONSHIPS
The main simplifications are that the animal is assumed to make use of steady-state flow patterns only, that lift is produced at right angles to the direction of the relative wind, that the stroke plane is horizontal with no tilt, that the induced wind can be disregarded because it is relatively small and, finally, that the wing movements are sinusoidal and of similar shape and the same duration for both the morphological
Flight fitness in hovering animals

dr
171
Fig. 2. Diagram of the wing parameters (A) and the movements during hovering (B).
upstroke and downstroke. As we shall see (p. 192), these assumptions lead to values for the average lift coefficients CL needed to remain airborne which are slightly higher than in the more complete treatment (Weis-Fogh, 1972). As to the specific aerodynamic power PJ, the estimates are too small by a factor of about 2 but, as we shall also see (p. 195), it is possible in a simple manner to make appropriate corrections. The instantaneous aerodynamic force on a wing element F(t, r) of chord length dr) and its vertical (lift) and horizontal (drag) components are shown in Fig. 1 A. In Fig. 1 and in the text, any quantity X which is a function of time t and distance r from the wing hinge, or fulcrum, is written as X(t, r). Appendix 1 gives a list of symbols and constants used. The justification for the simplifications is that the majority of insects, birds and bats have been found to tilt the long axis of the body towards the vertical (Fig. 1B) when hovering, so that the wings beat in an almost horizontal plane, usually symmetrically about the average positional angle y of 900 (Fig. 1C).
(a) Coefficient of lift
Fig. 2 A shows how the outline of a wing of total length R, and in particular the wing chord dj), varies with the distance r. In practice I have found that most wing
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TORKEL WEIS-FOGH
contours can be described quite faithfully by means of a simple mathematical functionl whether the moving wing is morphologically two-winged as in Lepidoptera and Hymenoptera or consists of only one part as in Diptera. It should be stressed here that there are hovering insects which make use of two pairs of wings which beat out of phase, as is the case in Odonata and Neuroptera, and these insects are not included in the present analysis. The angular movement of the long axis of the wing in the horizontal stroke plane is 7(0 = y + W>sin(2nnt), (1) where y is the average angle, <f> the stroke angle, n the wing-stroke frequency (number of complete stroke cycles per unit time) and t is time. The angular velocity and acceleration are then
dyjdt = 77710 cos {zrnit), (2)
d*yldt = -2nW<f)sin(z7mt).
(3)
The instantaneous force on a wing section at distance r from the fulcrum depends on the square of the instantaneous velocity of the section (v(t, r))2, on the force coefficient, and on the area of the segment of width dr (see Fig. 2 B). In the case of the instantaneous lift produced by the section, we then have dL(t, r) = \px CL(t, r) x area x velocity2. (4) Thus (5) dUt, r) = \pxCL(t, r)xc{r)drxr\dyjdtf, where p is the mass density of air and GL(t, r) the instantaneous coefficient of lift at distance r. This means that (6) dUt, T) = i/OT^V2 * CL(t, ')x c(r)ridr x cosi(2nnt). The four terms will be treated in turn. The first is a constant. The second (CL(t, r)) will be taken as constant in this treatment for the following reasons. In large insects and hummingbirds the wing is twisted linearly with respect to r so that succeeding sections have the same geometrical angle of attack (Jensen, 1956; Hertel, 1966). In small insects like DrosopMla the wing is rotated as a whole plate but the lift coefficient hardly varies at all with the angles of attack actually applied by that insect (Vogel, 1967a). In both cases the angle of attack is set almost instantaneously at the beginning of each half-stroke and remains constant so that it is justifiable to consider the average coefficient of lift CL as a constant, as previously discussed (Weis-Fogh, 1972). Of course this need not always be the case, but it is a reasonable approximation to begin with. It also has the advantage that one can use steady-state lift/drag diagrams for fixed wings of known shape and aspect ratio later in the analysis. The third term in equation (6) is the derivative with respect to r of the second moment S of the total wing area of all wings about the fulcra; it depends only on how the area varies with the distance r from the base, i.e. it is a function of the contour of the wing. Since many biologists are unfamiliar with this expression it is treated in Appendix 2. The general expression is S = <rcR\ (7) where a is a shape factor characteristic for the particular wing shape. It can be found by graphical integration based on actual wings or from the mathematical function by
173
Table 1. Shape factor afor the second moment of the area of two whole functional wings (or two pairs of wings) (eq. 7), shape factor rfor the third moment of two whole wings (eq. 50), together with the radius of gyration rg, the chord length ca at distance r^ the factor in eq. (11 b)for the calculation of Reynolds number for the wings shapes considered, and the factor in eq. (35)/or calculating A, the ratio stroke-arc I wing-chord.
(a) (c)
Parabola '. tectangle
hape...
Half ellipse
-
Triangle, base c
Triangle, apex
^^
i f
o-4ic 3-3 O-995 o-yic io-o 0-995
Tot
r,
c, Factor in eq. (116)

Factor in eq- (35)
087c 8-7
"575
\R
R/VA
1 I
I-OOC
0-690 6-6 0693
n-5
O-S77
means of which the outline of the wing is described. The chord of the wing c is then defined in each single case, as is explained in Appendix 2 and Table 1, where some shape factors are listed. It should be noted that the third term is a function only of r and not of t. The fourth and final term in equation (6) is a function of time / and not of distance r, and we therefore have, for a complete quarter-stroke, that the average lift L is
L=
X I*'
J r-0
c(r)rldrx [ ~
J t-0
cosi(27mt)dt.
(8)
A quarter-stroke is the time which it takes for the wing to move from an extreme position and until it reaches the middle of its stroke path. We have already dealt with the first integral, which depends only on the wing shape and is equal to S. The second definite integral equals $. Since the movements and forces are assumed to be symmetrical about the middle position (harmonic motion) the average lift over each quarter-stroke must equal the weight G of the animal so that the average lift coefficient is (9) (b) Reynolds number (Re) For a given aerofoil the coefficients of lift and drag depend not only on the angle of attack but also on Reynolds number (Re). This is particularly so in tiny insects, where it may be as low as 10 or even smaller (cf. Horridge, 1956). The task is to find a simple but representative expression. The general expression is
fl V '
V
'
where /* is the viscosity of the air and the ratio (i\p is the so-called kinematic viscosity v. In all systems of measurement, v has the dimension of (length)2/time. At 20 C and
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TORKEL W E I S - F O G H
normal atmospheric pressure v is o-i4cma/sec in the units used here. In ordinarf fixed-wing airplanes the representative velocity is the airspeed and the length is some representative wing chord. In our case we much make a specific choice. During hovering 80% of the lift is produced during the middle part of a half-stroke, where the average value of cos(27rcif) in equation (2) is 0-89. As to distance from the fulcrum r, we will choose the radius of gyration rg of the second moment of the area of the chosen wing shape (whether rg is determined graphically or analytically) and use the chord length at that distance, cg. Some useful values of rg and cg are included in Table 1. We then have = Since rg and cg are functions of R and c, a practical expression for Reynolds number is (Re) = factor x cRnfi, ( I r *) where the factors relating to the wing shapes used in this study are also included in Table 1. (c) Drag, aerodynamic torque and power As a corollary to equation (5), the instantaneous drag D(t, r) experienced by a wing section is dD(t, r) = ip x CD(t, r) x c(r)r*dr x (dyjdtf. (12) The bending moment Qa about the fulcrum, which this gives rise to, is the product of the drag and its distance r from the wing base, thus dQaV, r) = lp* CD(t, r) x c(rydr x (dyldt)*. (13) Since we have given reasons for using the value of the lift coefficient as averaged over both time and distance in this particular type of flight, we must do the same for the conjugated drag coefficient so that the second term is a constant CD. The third term is the derivative with respect to r of the third moment of the area of all wings about the fulcra dT(r). The instantaneous aerodynamic bending moment of the wings which the thorax must act against is therefore
Qa{t) = \pCD x I]]**)**
{^j\
(14)
The definite integral is the third moment T of the wing area about the fulcrum. It is not normally encountered in engineering and it is therefore treated in Appendix 2, from which it is seen that T = TCR*, where r is the appropriate shape factor (cf. Table 1). The bending moment then varies with time as Qa(t) = yC^cR^n^n^cos^znnt). (15) The aerodynamic work done by the thorax on the air during a quarter-stroke is
which, when combined with equation (14), gives
3 J 1 - 0 cos (znnt)dt.
f t-
in
(17)

during a complete wing-stroke cycle consisting of 4 quarter-strokes is
175
Pince the value of the definite integral is 1/(37771), the total work done by the wings
(18) Wa = $pCDTc&n*n*p. During steady-state hovering the average aerodynamic power which the thorax must produce in order for the animal to remain airborne is therefore (19) ?a = nWa = $pCDTC&nW<p. In addition to the quantities already discussed in equation (18) and (19), we must find CD in order to apply the equations. In our case, and since we base the analysis on steady-state flow, this is done by using experimental lift/drag diagrams of known wings of suitable form and aspect ratio. Ideally one should use the actual wings of the insect in question, but this is only possible in a few cases at present. The chosen diagram is stated in Table 4 for each species considered. The choice is based upon wing shape, wing profile, Reynolds number and, of course, availability. When CL and (Re) are known, CD then follows, but apart from locust wings (Jensen, 1956), the other estimates suffer from the shortcoming that the lift/drag diagrams were obtained in a linear air stream of uniform velocity. The rotational components, the variation in speed from base to tip and the effects that this may have on details of the flow are therefore disregarded. Future experiments may remedy this shortcoming but the effect is not likely to be serious for a first approximation as attempted in this paper. However, there may be exceptions for some species as pointed out in the Discussion.
WING INERTIA AND DYNAMIC EFFICIENCY
If / is the moment of inertia of the mass of all wings about the fulcra, the acceleration of the wings gives rise to an inertial bending moment Qim According to equation (3) it is Spy QJt) = l- = -2/7rV0sin(27m*). (20) During harmonic oscillations Q will then vary linearly with the positional angle y and be maximum at the extreme wing positions and zero at the middle of the stroke. Numerically the maximum value is (21) |?i|max = 2lnW<l>. According to equation (15) the aerodynamic moment is maximum when the wings pass their middle position, zero at the extreme positions, and |?| max = kpCDTcRhT^(j>\ (22) In order to calculate the total work done by the thorax when both inertial and aerodynamic bending moments are taken into account, we must integrate Q^+f, over a complete wing stroke. This was discussed in detail in a previous paper on the basis of some actual examples (Weis-Fogh, 1972). The present account is easier to follow if the reader is familiar with the approach given in that paper, and it should be emphasized that the following derivations are valid only if elastic torques inside the thorax or in the wings themselves are disregarded. The aim is to deduce a simple analytical expression for the relative importance of inertial forces as compared with aerodynamic forces in the absence of elastic defor-
176
I
TORKEL WEIS-FOGH
I
20
30 40 Distance from fulcrum (mm)
50
60
Fig. 3. The distribution of the wing weight as a function of the distance from the fulcrum in a standard Sckutocerca gregaria. (A) One forewing (1) and (B) one hindwing (2). (C) is a model of the mass distribution and applies to insects in general.
mations and storage of energy. This is most easily achieved if we use the ratio N between the maximum inertial and aerodynamic moments in equations (21) and (22), according to which
It is now possible to find an expression for the dynamic efficiency 7 7 of the system, where T\ is the ratio between the total aerodynamic work and the combined inertial plus aerodynamic work. Again disregarding elastic deformations, V = However, we must first find a value for /. (a) Mass moment of inertia In Table 4, / is the moment of inertia of all the moving wings about the fulcra, but it is simplest to consider a single fully unfolded wing. The mass of a narrow transverse wing section at distance r is pc(r)d(r)dr, where p is the average mass density of the wing material, d(r) the average thickness, dr the width of the section and c(r) the chord. I = p[~Rc{T)d{T)r*dT. (25)
(24)
In practice, I is usually found by cutting the wing into strips normal to the long axis, estimating the mass of each section by weighing and finding the sum of the products of mass times distance squared. In insects, however, I have found that an easier procedure is often possible and sufficiently accurate for our purpose.

Table 2. Moment of inertia I of the mass of one fully unfolded fresh vring about the main vmg hinge in a standard Schistocerca gregaria
(Ten old and 11 young adults of both sexes. The experimental results are compared with those calculated from equation (27). There is no significant difference.) One forewing Length R (mm) Weight (mgf) /(gf cm sec1): Experimental Calculated 57-1 -i 29-5 0-7 io~* (i - 7io-o4) 10-4 (1-63005) One hind wing S3-i -2 23-610-5
io~* (I-IOO-O3)
177
10-4 (1-1310-03)
Locust wings. The distribution of mass in fresh wings of Schistocerca gregaria is seen in Fig. 3 A and B. The insects were of different sizes but all results were recalculated to a locust of average size, a standard Schistocerca, by multiplying lengths with the ratio of the length indices, ///s, and weights by the ratio of the volume indices, v/v,, where /, and vs are the standard indices for length and volume and / and v those of the individual insects used (Weis-Fogh, 1952). The standard deviations of the corrected figures from 21 males and females of different adult ages are shown as vertical bars. When a straight line is drawn from the wing tip and adjusted to follow the curves as closely as possible, it is seen that the mass tends to decrease linearly with distance r from the fulcrum both in the broad fan-shaped hindwings and in the stiff slender forewings. The deviations occur mainly towards the base, where r is small and the effect negligible. Consequently we can calculate / as being equal to that of a homogenous triangular plate of uniform thickness d and which is rotated about its base line c (Fig. 3 C). In this case c(r) = c{R r)/R, and a strip of width dr has the mass pdc(R r)/R x dr. We then have
{y
r-0
( 26 )
(27)
Since the total mass of the plate is M = \pdcR,

1 = *M#.
This simple expression gives values for locust wings which are accurate within the limits of the experimental estimates based upon weights and distances, as is seen in Table 2. Other insects. It is often difficult to cut and weigh small insect wings, and for quick estimates it is obviously advantageous to use an expression like equation (27). Is it applicable to other insects? In order to answer this I have calculated the moment of inertia by means of equation (27) for the insects in which Sotavalta (1952, 1954) found / by weighings. His results are of course not as accurate as those for the locust because the wings are small and only a few strips could be cut. Nevertheless, the conformity between the calculated and the empirical results is good, as is seen from Table 3, in particular when the sources of error are taken into account. In fact, if accurate measurements are not readily available, the formula provides values which are entirely adequate for the present treatment. For flying insects considered as a group, the formula
ia
EXB
59
178
TORKEL WEIS-FOGH Table 3. The moment of inertia of the wing mass about the wing hinge in different insects
(The deviations of the values calculated from equation (27) and those found by Sotavalta (1952, 1954) are given as a percentage.) No. of examples 3 3 8 Genera considered (deviation in parentheses) Sympetrum ( + 22), Aetkna ( 35) Mdolontha (o), AmpkimaUon (o), Cerambydd ( + 5) Agrotii ( 11), Ampkitrota ( 5), Sideridu ( 20), Amathet ( 21), Poecilocampa ( 8), Macroglossum ( + 17), Pieris(-i9). Apis (17), Bombus ( 16), Vespa (14), Opkion (-22) Tipula (-36), Trichocera (-15), Theobaldia (-28), Eristalis ( + 9), Calliphora ( + 7). Average deviation
-i6
Insect order Odonata Coleoptera Lepidoptera
+2
II
Hymenoptera Diptera
5 5
-19
is accurate and it must reflect some structural principle of general significance, to be discussed in another context (Weis-Fogh, in preparation). Effect of wing mutilation. In order to discuss some results reported in the literature (Danzer, 1956) we must deduce how / alters when the wings are mutilated. We assume that / = ^MR2. Let the moment of inertia of the mutilated wing be Im. It can now be calculated by means of the theorem of parallel axes. In Fig. 4A let Mx be the mass of the outer cut section whose centre of gravity is distance r from the fulcrum. The moment of inertia of the cut section about the fulcrum is now Ix = I0+Mxr*, (28) where / is the moment of inertia of the section about its own centre of gravity. It is 70 = ^MjA:2, where x is the distance from the wing tip to the mutilation cut m. Let the relative distance from the tip be so that = o at the tip and = 1 at the base. We then have that * = R, Mx = ME? and r = R-^2Ji) (see Fig. 4A). Thus Ix = For the mutilated wing 7m = / - / x = Mi?*(i-/()). (30) The numerical value of the functions/() and (1 /()) are plotted in Fig. 4B. These curves are of general validity provided that the wing mass decreases linearly with r, i.e. provided that equation (27) is a sufficiently good approximation for the particular enquiry.
(b) Dynamic efficiency - (29)
As discussed earlier (Weis-Fogh, 1972), we shall not count work which is absorbed by the working muscles in the thorax by their braking action, the so-called negative muscular work, but only the positive work imparted by the thorax to the moving wings. In order to do this our task is to integrate the function of the combined aerodynamic and inertdal bending moments Q^^ (7) with respect to y when the wings move from one extreme to the other, i.e. over one complete half-stroke. This is sufficient since the

Fulcrum
179
1
*
1- - " T
10
0-8 -
i-AD \
/AD
0-8
I 0-6
- 0-6
\
B 0-4 -t
i i
\
\
r
0-4
0-2 -
- 0-2
.'''
T-L
0-2
0-4 0-6 0-8 Relative distance from tip,
10
Fig. 4. The effect on the moment of inertia / of mutilating the wing of an insect by means of a tranverse cut (m), as explained in the text and in equation (30).
two half-strokes are similar with respect to work. More precisely, we want to find the ratio between the following two integrals, eventually expressed in terms of the ratio N:
QM<ir.
(3 1 . 32)
An exact solution is lengthy and tedious to use. The procedure adopted here is neither exact nor elegant but it leads to a simple approximation which is accurate to within a few per cent, which is more than adequate for our purpose. Since we are dealing with ratios rather than absolute figures, let the angular movement represent 1 angular unit so that the wing moves from y = o to y = 1 during the upstroke in Fig. 5. Let the maximum aerodynamic moment in equation (22) be represented by + 1 unit. The maximum inertial moment in equation (21) is then +N units at the beginning of the upstroke and N units at the end. While Qi varies linearly, Qa varies according to the thick curve ABC. The combined curve Qfa+f) is represented by the thin curve from + Af to E'. It is the area between the latter and the base line AE' which represents the total positive work during one half-stroke. If the aerodynamic torque also varied linearly from minimum to maximum and from maximum to minimum, the work during the first quarter-stroke would correspond to the area of triangle ABD and be \ work units; during the second quarter it would be epresented by DBC and also amount to . In this case the combined work J ^
1
i8o
+3 +N +2
TORKEL WEIS-FOGH
\\
I-.-2 -
\\
\\
X
\\
\
\ \ \ \ \ \ \
N. \
\\ -N
7 = 0-5 JW=l Positional angle y (relativo units) Fig. S- The work diagram of a hovering animal and the method of integrating the total work during one half-stroke, as explained in the text.
-3
would be represented by the area of the quadrangle ANBE, which is -NJ4{N+i)) work units. However, we have to add the two dotted areas above the lines NB and BE' respectively. From equation (17) we already know the exact value of the aerodynamic work during one quarter-stroke. In our case the area is \ work units so that it is ^ unit larger than the triangular area during the first quarter. During the second quarter the area to be added is approximately equal to the densely dotted area EBE", leaving out the small horizontally hatched area EE'E'. The dotted area is equal to the ^ unit times the ratio DE/DC, which is i/(iV+1). We then have for one half-stroke that N 1
24(iV+i) '
For large values of N the error is insignificant and for very small values it is also unimportant, amounting to 0-2 % for N = o, i.e. when the inertia forces are zero. The error may rise to a few per cent at intermediate values; this does not matter here. Since Wa amounts to $ work units per half-stroke, the dynamic efficiency is V = Wa
(34)
181
Fig. 6. The relationship between the dynamic efficiency rj (in the absence of elastic forces) and the ratio N between the maximum inertial and aerodynamic bending moments in equation (33).
N is easily found from equation (23) and the relationship between 7 7 and N is plotted in Fig. 6, from which the dynamic efficiency can be read. When the inertial forces are vanishingly small and N = o, the efficiency is of course i-oo. When the two bending moments are equally large it is 0-85, but for N = 5, which is quite common, V is 0-37.
EXPERIMENTAL METHODS AND MATERIAL
Most of the flight data derive from critical examination of the literature, but it was essential to obtain new information concerning both large and very small insects. (a) Syrphinae hovering in nature The true hover-flies belong to the subfamily Syrphinae within the large family of Syrphidae. So far, the best studied genera have been Volucella and Eristalis, which represent two other subfamilies; they hover in a way similar to that of wasps and bees, i.e. in accordance with our main assumptions (see the definition of 'normal hovering' in the next section). However, the Syrphinae fly forwards, sidewards and hover with the body axis horizontal and with the wings oscillating in a non-horizontal plane and through a surprisingly small stroke angle. Obviously, they are exceptions to the general rule; the same applies to the Odonata (damselfliesand dragonflies). While I have been unable to study hovering dragonflies in nature, I have recently succeeded in recording the wing-stroke frequency of Syrphinae of the genera Syrpkus, Sphaerophoria and Platychirus together with the body weights and wing parameters of the same individuals. The animals hovered in front of flowers or between vegetation where there was no wind on calm summer days at Tibirke in Denmark. The flight tone and therefore the frequency was picked up by means of a Briiel & Kjaer 25 mm Condensor Microphone
182
TORKEL WEIS-FOGH
Type 4145, with battery-driven Pre-amplifier Type 2619 and Power-supply Typl 2804. The signals were recorded on a Nagra IV-S battery-driven tape-recorder supplied with a special noise-reduction system which ensured flat response characteristics between 100 and 970 Hz. The steep cut-off at either end corresponded to a reduction of 20 dB at 30 and 2000 Hz respectively. This was essential in order to prevent interference from other sound sources. The individuals were narcotized instantaneously while hovering by means of a strong jet from a Sparklets CO2-spray filled with chloroform and caught in a small net. The weights of the body and the wings were determined by means of an electronic microbalance immediately on return to the laboratory. The insects were identified according to Coe (1953). (b) Free hovering in the laboratory Large dung beetles. The flight of medium-sized and of small lamellicorn beetles like Melolontha vulgaris (cockchafer), Rhizotrogus (now AmphimaUon) and Cetoma aurata (rose chafer) have been studied both in nature and in the laboratory (Sotavalta, 1947, 1952), but since the papers by Osborne (1951) and Bennett (1966, 1970), the aerodynamic mechanisms involved have been in dispute. In particular, Magnan's (1934) data for the large stag beetle Lucanus cervus have been taken as evidence for substantial contributions by non-steady flow. Thus Osborne (1951) found an average lift coefficient CL of more than four. The beetle weighed 2-6 gf and had a' wing load' of 3*2 kgf/ ma. I could not obtain this species but Professor J. W. S. Pringle, Oxford, brought some large elephant-dung beetles of the genus Heliocopris back from Kenya. He kindly measured the frequency acoustically during near-hovering, free flight and also gave me weights (8-13 gf) and wing dimensions; the wing load was 3-4 kgf/ma and therefore comparable to that of the stag beetle. As in other lamellicorn beetles, the stroke angle <j> is about 1800. Large hawk moths. Dr P. J. Wilkin, Brunei University, has recently shown me (personal communication) recordings of wind-tunnel experiments with the Florida tobacco hornworm moth, Manduca sexta (Johannson), which was flying suspended almost horizontally from an aerodynamic balance. His results indicate that, under certain conditions, these insects can produce lift coefficients considerably in excess of steady-state values, and he seems to have confirmed this in model experiments. The question is how and to what extent freely flying sphingid moths make use of unusual aerodynamics - for instance, when feeding normally like hummingbirds in front of a flower. Caterpillars of Manduca sexta (Sphingidae, subfamily Acherontinae; earlier name Protoparce sexta Johannson) were grown in the laboratory on a semi-synthetic diet (Hoffman, Lawson & Yamamoto, 1966). Adults were isolated and hovered spontaneously in front of an artificial flower but only shortly after ' dusk' and in subdued light. The flower was scented with the synthetic attractant isoamyl salicylate and the feeding solution contained 10% sucrose in water. The temperature was 26 C and the relative humidity 90%. The flight chamber measured 71 x 71 x 62 cm3, and when the moth was hovering in the correct place near the centre of the almost dark cage two crossed light-beams of low intensity were interrupted by the insect. This blanked two C.D.S. phototransistors which triggered a time-delay circuit which then started a sound-insulated high-speed 16 mm motion camera (Hitachi HIMAC Type 16 HM, rotating-prism type, with speed control unit HM 502 and time switch to prevent
183
iverride). It also triggered two 1000 W quartz-iodine lamps, and the timing was such that the lamps did not light up until the film had reached a sufficient speed, i.e. about 3000 frames/sec. This elaborate arrangement was essential because Manduca sexta is a nocturnal insect which often makes a dive when struck by intense light. However, since this reaction does not take effect for a few wing beats, normal hovering as well as aerobatics and fast forward flight could be studied, at least in brief sequences. Small chalcid wasp. The flight of very small insects is virtually unknown (Horridge, 1956). Pupae of the parasitic chalcid wasp Encarsiaformosa Gahan (family Eulophidae, subfamily Aphelinidae) were obtained from a commercial dealer; this species is used for biological control of the greenhouse white-fly, Trialeurodes vaporariorum Westwood (see Speyer, 1927; Burnett, 1949; Parr, 1968). The tiny adults (wing length 0-05-0-07 cm) both fly actively and make strong jumps. The procedure therefore was to take a high-speed film (as above) whenever the wasps became particularly active, as could be brought about by gently tapping the flight chamber. The plastic chamber was 5-4 cm high, 3-5 cm wide and 1 -6 cm deep, so that the insectsflyingnear the centre were within the range of focal depth (2 mm) and aerodynamically almost free of wall effects. The insects are diurnal and are attracted to light. A 50 W fibre-optical vertical light beam was constantly on and tended to concentrate them in the vertical centre axis of the chamber, while three 150 W quartz-iodine lamps were triggered simultaneously with the camera (6-8000 frames/sec); they did not appear to interfere with flight. The lens was a Carl Zeiss 63 mm Luminar,/1:4'5, used at low focal opening, and the magnification on thefilmwas one to four times. The temperature was 26-27 C and the air almost saturated with water vapour. It was usually easy to distinguish between jumps, flight initiated by a jump, and free 'unaided' flight. Only the latter scenes are used here.
RESULTS
As in the previous study (Weis-Fogh, 1972), I shall use the technical force-lengthtime system, i.e. gram force-centimetre-second or gf-cm-s system, but the relevant SI units and conversion factors are given in Appendix 1.
(a) Normal hovering
All birds and bats, and many insects, make use of fast forward flight for transport over long distances; locusts and other migrating species are good examples. This type of flight is now reasonably well understood in birds and insects on the basis of ordinary aerofoil action (Weis-Fogh & Jensen, 1956; Jensen, 1956; Weis-Fogh, 1956; Pennycuick, 1968 a, 1969). However, many small birds and bats and most insects also practise slow forward flight and hovering on the spot when they forage, approach the nest, perform sexual displays or explore the surroundings. The explanation is not only that this, the most strenuous form of flight, is less demanding in small animals than in large animals (because of the surface/weight ratio), but presumably also that the conquest by means of active flight of the small niches of the aerial biosphere has opened up an endless variety of ecologically different niches to which the response has been intense speciation, best exemplified by the 750000 known species of winged insects. Hovering flight is therefore of general significance, particularly in insects. The main problems are whether it is based on a common mechanism or not, and what the main principle is.
184
TORKEL WEIS-FOGH
(b)
(c)
Fig. 7. Typical body and wing postures in some hovering animals, as drawn from flash photographs of (a) the hummingbird ArchUochuz colubru (Greenewalt, i960), (6) the sphingid moth Deilephila elpenor (Nachtigall, 1969), (c) the sphingid moth Manduca sexta during normal hovering (present study), (d) a bumble-bee Bombus sp. before landing on a flower (Schmidt, i960), (e) Manduca sexta during a quick manoeuvre (present study), and (J) the cockchafer Melolontha vulgaru during vertical take-off (Lane, 1955).
185
We have already seen that the hovering of hummingbirds and of Drosophila involves the same type of wing movements and can be understood on the basis of steady-state aerodynamics, although non-steady periods and effects undoubtedly exist and may play a significant role under some conditions (Weis-Fogh, 1972). Both types of animal perform what I shall call normal hovering. It can be defined as (a) active flight on the spot in still air by means of wings which are moved (b) through a large stroke angle and (c) approximately in a horizontal plane, while (d) the long axis of the body is strongly inclined relative to the horizontal, sometimes almost to a vertical position. Fig. 7 shows some examples of normal hovering. It has been drawn from flash photographs of freely flying animals. The first question is how common normal hovering actually is. All the animals listed in Tables 4 and 5 are known to hover, and if the name appears without an asterisk the species uses normal hovering. The examples have been greatly extended by direct observations of free flight in nature which I have made during the past few years. In the case of nocturnal species a high-pressure mercury-vapour lamp was used (Philips HP 125 W, 4000-7200 A). Normal hovering was observed in many species of Isoptera in East Africa (swarming termites), Plecoptera (stone-flies), Thysanoptera (thrips), Hemiptera (leaf-hoppers, aphids, backswimmers, may bugs), and Neuroptera (lacewings). The Coleoptera or beetles deserve more comment. Although the elytra (forewings) are usually spread out and flap to some small extent, they contribute next to nothing during hovering flight (Nachtigall, 1964) and the hind wings can be considered as the sole movers, beating in a horizontal plane (cf. cockchafer in Fig. 7/) and sweeping through a large angle of about 1800 on each side. However, because the body is broad, the two wings do not come into]close contact. Normal hovering is perhaps more easily studied in Coleoptera than in any other group except the large crane-flies (Tipulidae), and most species or groups make use of it, the tiger beetles (Cicindelidae) being possible exceptions. Among Lepidoptera the moths generally appear to move their wings in accordance with the common pattern. This is indicated by my direct observations and also appears from flash photographs taken in nature (Fig. 76, Deilephila elpenor) and it was proved by the slow-motion film9 of the sphingid moth Manduca sexta hovering in the laboratory (Figs. 8, 9). In both cases the moth was photographed vertically from above. Fig. 8 shows one complete wing stroke during normal hovering on the spot. It often happens that the wings of one side are leading slightly, relative to those of the other, and in this case the left wings were leading. The morphological underside of the wings i9 black. The sequence started when the left wings began the morphological downstroke (Fig. 8 a). The film speed was 3000 frames/sec, increasing to 3900 at the end of the stroke (Fig. 8p). Only every 10th frame was traced. From the apparent length of the body axis it is seen that the body is inclined about 45 relative to the horizontal. This scene, and films taken horizontally from the side, show that the wings beat almost horizontally like those of hovering hummingbirds, that they are twisted in a similar way and that upstroke and downstroke are of equal duration (cf. Weis-Fogh, 1972), the tilt /? of the stroke plane being about io. It is apparent from the films that the fore- and hindwings move together and that a cleft between them does not normally open, although this happens during certain brisk manoevres. Furthermore, during normal hovering the acceleration of the wings does not lead to any significant elastic
i86
TORKEL WEIS-FOGH
(0
(m)
()
0
5 cm
V ()
(0
(p)
Fig. 8. Normal hovering of the sphingid moth Mcmduca sexta as traced from every ioth frame in a slow-motion film taken vertically from above at 3000 (start) to 3900 (end of stroke) frames/sec. The undersides of the wings are drawn in black.
bending of the long axis of the wing (Figs. 8a-p; jc) but, again, substantial bending can occur during quick escape reactions (Fig. je). We are not concerned with the latter situations here; the main point is that the elastic storage of energy is confined to thoracic structures during normal hovering as well as during fast forward flight, as is apparent from other films. Fig. 9 is an example of part of a complicated manoeuvre where the moth started from normal hovering in front of theflower,but after the light and camera had been on for a short time it began to move backwards with vertical body axis (Fig. ga; 3000 frames/ sec) and during the stroke the axis tilted backwards and the speed was also increased in the backwards direction. The morphological underside (black) of the wings is therefore seen during the main part of the stroke, whereas the upper side would predominate during fast forward flight, underlining the supreme manoeuvrability of the insect as well as the general principles of flight. On occasions like this the manoeuvre even resulted in a backwards somersault. The flight of true butterflies (Papilionidae) is different from that of sphingid moths, partly because the wing surfaces often come into direct contact with each other at the top of the upstroke (a ' clap') and partly because it looks as if the angular movements deviate appreciably from the sinusoidal case. They have therefore been marked by a double asterisk () and need to be studied separately in another context. The Mecoptera (scorpion flies), Trichoptera (caddis flies) and the large group of Hymenoptera (sawflies, swarming ants, wasps and bees) all appear to use normal hovering, at least as the main principle. It is well known that the honey-bee (Apis
T87
M
00 CO (?) (A)
(0
("0
' '
()
IMMI 0 5cm
'
()
'
(P)
Fig. 9. Same as Fig. 8 but during the beginning of a backwards somersault, a explained in the text.
mellifica) can change the inclination of the stroke plane and that this is almost horizontal when the insect is hovering in front of aflower(Neuhaus & Wohlgemuth, i960; Nachtigall, 1969). It is more difficult to observe this in the bumble-bee (Bombus) because of the transparent and relatively small wings. However, my own observations and Fig. y(d) indicate that they also use normal hovering. This was easy to observe with certainty in the large carpenter bees of the genus Xylocopa, which have deeply stained wings (X. violacea in Italy, 1972; and an unidentified species in Kenya, 1972). It may have been thought that the very small chalcid wasp Encarsia formosa would represent an exception but this is not the case, as is apparent from Fig. 14 and the description later. The reason for the triple asterisk (***) against its name in Tables 4 and 5 is that in every single wing stroke the two pairs of wings come close together in a distinct dorsal clap before the morphological downstroke starts. The significance of this will be analysed in the Discussion (p. 214), but the same phenomenon appears to be present in Drosophila virilis according to Vogel (1965, fig. Ill, 1), who did not comment upon it however. It has therefore been similarly marked. The large and diverse group of truly two-winged insects Diptera presents special problems. Whereas crane-flies, mosquitoes and other Nematocera as well as many large Brachycera and Cyclorrhapha undoubtedly use normal hovering in most cases (this applies for instance to Eristalis, Volucella and probably Calliphora), the dorsal clap seen in Encarsia and Drosophila probably occurs in many other small flies and insects from different orders. In addition, the true hover-flies, the Syrphinae, perform steady hovering flight according to principles which are unknown so far and which
i88
TORKEL WEIS-FOGH
(b)
(c)
00 Fig. 10. Examples of wing shape and applied contour functions in (a) the hummingbird AmaziUafimbriatafltwiatUU, (b) thefruit-flyDrosophtlafimebris, (c) the hover-fly Syrphus ribem, (d) the hornet wasp Vespa crabro, () the sphingid moth Manduca sexta, and (/) the cabbagewhite butterfly Pieris brasticae. The hatched areas are discussed in the text.
also seem to apply to large dragonflies, Odonata, as already pointed out. Their flight will therefore be treated separately. Finally, some small bats make regular use of hovering, and although the main movements are consistent with the normal pattern the wings are somewhat flexed during the morphological upstroke (Eisentraut, 1936; Norberg, 1970), as is also the case in small passerine birds (Zimmer, 1943; Brown, 1951; Greenewalt, i960). A single asterisk (*) is therefore put against Plecotus auritus in Tables 4 and 5. The term normal hovering is then well justified, the most important known exceptions being Papilionidea, Syrphinae and Odonata.

(b) Wing shape and contour function
189
During hovering it is the outer two-thirds of the wings which matter. At a first glance it may seem difficult to find a simple geometrical function which describes the wing contour in sufficient detail for a realistic estimate of the second and third moment of the area, S and T. However, it is not difficult in practice, and in most cases the wing(s) can be described by means of a semi-ellipse where the wing length R corresponds to half the major axis and the minor axis is adjusted to give the best fit for the outer part of the wing; this is the value for the adopted wing chord c in Table 4. The procedure is illustrated in Fig. 10 for both two-winged and four-winged animals. In some cases the major axis is tilted by a few degrees relative to the line from the fulcrum to the tip in order to give the best fit, as is seen in the case of Syrpkus (Fig. 10c). This does not introduce any significant error. It may be thought that the hatched areas between the wing contour and the ellipse justify a substantial correction, but this is not the case. If, for example, we consider a wing in which the inner part is constricted, leaving a large hatched triangular area, as in Drosophila and Syrphus, with a base equal to \c and a height of \R, S will be overestimated by 3 % and T by only 1-2%. The lift coefficient and the aerodynamic power will then be underestimated by similarly small amounts and this is far too little to worry about in the present approach. The poorest fit is undoubtedly that of Pieris in Fig. io(/) because the outer areas count most. However, the deviation is not large enough to be of real concern since much larger errors are introduced because our basic assumptions do not seem to apply to this group. In quantitative terms, the fit for the hummingbird AmaziUa and for Manduca is almost perfect, taking into account that the contour does not remain really constant in any animal during actual flight. The special problem of Encarsiaformosa will be discussed in connexion with Fig. 13.
(c) Flight data: Table 4 Only the most reliable data (or free flight have been accepted and only if they belong to one set for each specimen included in the list. The data for Drosophila virilis are an exception since they derive mainly from Vogel's wind-tunnel studies, as discussed elsewhere (Weis-Fogh, 1972). As to Encarsiaformosa, the weight G is the average from some ten individuals and the first example refers to a particular flight of a particularly large specimen, whereas the second entry represents average values. Apart from body and wing dimensions, the most critical parameter is the wing-stroke frequency n because it varies greatly with size and it enters into the equations in the second or third power. However, in the selected examples the values can be considered to be accurate. The same does not apply to the stroke angle <fi, but it cannot normally amount to more than 1800 = n = 3-14 rad, and <j> is usually about 1200 = 2-09 rad in both large and small hovering animals. The ratio A between the stroke arc and the wing chord deserves special attention because, when an ordinary wing starts to move, it takes 1-2 chord lengths for circulation F to build up, i.e. for full lift to be produced. Consequently, I have calculated the stroke-arc/wing-chord ratio A at the distance from the wing base corresponding to the radius of gyration rg for the second moment S of the wing area about the fulcrum
Table 4. Flaght data used in this d y &
T h e most reliable values measured during free hovering flight, slow forward flight or, in a few cases, in wind-tunnel experiments. T h e references are given in parenthesea. I n thia and the following table the numbera mean: (I) Sotavalta, 1952, (2) Sotavalta, 1954, (3) Vogel, 1966, (4) Weis-Fogh, 1956, (5) Weis-Fogh, 1961,(6) Weis-Fogh, 1972, (7) Hocking, 1953, (8) Neuhaua & Wohlgemuth, 1960, (9) Norberg, 1970, (10) Weis-Fogb, unpublished, (11) Pnngle, personal wmmunication, (12) Sotavalta, 1947, (13) Greenewalt, 1960, (14) Greenewalt, 1962, (15) present study, (16) L a s i d , Weathers & Bemtein, 1967, (17) Prandtl& Teitjem, 1934, (18) Jmsm, 1956, (19) Nachtigall, 1967, (20) Vogel, 19676, (21) Horridge, 1956, (22) Hertel, 1966, (23) L a a i e d , 1963, (24) Chadwick & Gilmour, IW, (25) Sotavalta & Laulajainen, 1961, (26) Thorn & Swart, 1940. The etroke-arc/wing-chord ratio h = ~ J c is , included as a baais for discuasion of non-steady aerodynamics. One asterink (+)meam that the species hover in a way somewhat different from the 'normal'. Two aster& (a*) indicate buttefiea and three aaterieks (**+) against Enunsia fonnosa and Drosuphila oiriliz mean that the upper wing surfacea may approach each other closely at the top of the etroke, here called 'clapping', although the wings beat horizontally and are moved basically aa a&&ned &the theory. WingMoment of Liftldras Wing stroke Stroke diagram, References Weight, ahape frequency, angle, Smk"a" references, onstrohe Wing-chord G aspect ratio 0 (9)
Bats: Plccotus m ' t u s *
9.0
Birds: hummingbirds, Tmhili
Arc-
colubTir
AmagiLia jimbriata j-71Niacilir
3'7 5-1 20'0
Palagma gigas Coleoptera: beetlea Melolonth0 *mis Amphimollon rolzcitiolt H e h a p n i sp. H. sp. Cctoltio muata Cerambycid species Lepidoptera: buttertliea & motha
0.60 0.29 7'8 I 2.8 0.46 0.14
pieriz ,+*+
P. brarsicac**
0.04 0.14 1-6 2'12
sphinx ligurtn' Manduca sexto M. swta Mocrogloura stcllatonon Amatha bicololago
1'12
0.28 0.09
Table 4 (cont.)
WingWing
Moment of inertia,
Weight,
I
(gf a n 8') x 10-I
Hymenoptera: wasps, been
(gf)
etroke Stroke ahape frequency angle, Stroharc (cf. n Wing-chord Table I) (sec-') (rad) A
v.wlgmir
v e s p a crabro
0.60
009
0.88 0.48
0'10
6I 2 . 7 15.8 13.1 0.85
Bombur t m e r t r i r B. lapidmiur Apir mefijica E w s i a fonnosa (Chalcid wasp)+++ E. f m s a (Chalcid wasp)+++
25 x 1025 x 10-
0.028
3X
0'010
6.6 0.06
1 0 '
0.001 0.15 0.056
2 . 7
078
I
0'002
IO-'
Table 5. Main results: c o e ~ t of s Iifi and drag, specific aerodynamic power and dynmtric e$%amq
The values of CLan not corrected b e c a w thin would make little difference. Aa to the epecif~c aerodynamic power both the uncorrected (equation 19) and the corrected values are given, and the valuer, for N and the dynamic efficiency q are both based on the corrected figures.) Specific aerodynamic power P: (per g f W y weight) (Eq. 19) 23)
L
c,
34) Dynamic
Simple
theory
corrected
Bats: P h h u mrthu Birds: hummingbirds, T d i

(7.0)
Metahlic rate during actual flight ( c a l h-l) per gf body weight
3.1
A T wkrb+ir Anraoiliofincbricrtapuoiotilu Patagona gigas Coleoptera: beetla MeIdontha vdgo+ir Amphimalla solrtlrialis
11.7 10.9 10.5
4'3 3.6
HH. ap.
sp.
11'3 9'4 18.5 20.2 15'7 8.2
3'4
Ceionio m a t o Certunbycid species Lepidoptem: butterfliee, moths Finir nap?
(0.4) 3'4 3.6 8.0 5'8
P. brassicae++ sphinx ligurtri Mcmduca sexta

(5.9) 7'3 6.2 5'3 6.9
M.rexto
6-2 5'9 3'1 5'0 6.3 9.2 7'2 I 6.4 r 1.5 9 " ' (9.7) (12-41
Macmglourcmrtellatmron Anratiru bicolorago Hymenoptera: wasps, beee Vupa crabro v.Oucpmir Bornbur im& B. lnpidmiur Apir mrUif;ca Eneorrin fonrtara (Chalcid wasp)+++ E. fancua (Chalcid wasp)+++
4.6
4 . 1
3'7 15.3 23'5 9'3 9'9
1'7 5-6 2.8

A = <t>rg\cg = factor x 4>Rjc.
193
(35)
. equation 7). We must of course use the chord length cQ at the same distance so that The factor in equation (35) is given in Table 1 for the different wing shapes. Except in some butterflies, A is usually larger than 2-5 and varies between 3 and 5 in most of the groups, which indicates that ordinary aerodynamics applies. However, in butterflies (Papilionidea) and in true hover-flies (Syrphijiae, Table 9) A is often as small as 1-4. Under these conditions one must expect non-stationaryflowto be of major importance. It should be noted that the small chalcid wasp has a relatively high value of about 4, but special mechanisms are definitely involved in this case. The reference numbers in the legend of Table 4 applies to Table 5 also.
(d) Gross results: Table 5
The weight of the animals ranges from 20 gf to 25 /Jgf and the wing length from 13 to 0-06 cm, the corresponding Reynolds numbers (Re) being 15000 and 15 respectively. It is possible to obtain lift coefficients approaching i*o during normal steadystate flow down to (Re) between 10 and 100, and the square-law relationship between force and velocity holds down to 10 (Thorn & Swart, 1940). It is therefore meaningful to apply the analysis to the entire size spectrum in the tables. However, in the range of (Re) between 10 and 100 the drag tends to be larger than the lift, for (Re) = 10 by a factor of about 3, and normal flight would be both difficult and expensive in energy. Above 100-200 the lift/drag ratio has improved sufficiently for normal principles to be operative. The major conclusion from Table 5 is that most hovering animals move their wings in accordance with steady-state principles and at Reynolds numbers well above the critical range, and so that the average lift coefficient CL is relatively small, seldom exceeding i-o. This applies to all the lamellicorn beetles hitherto considered exceptions (Osborne, 1951; Bennett, 1966, 1970) and to most wasps and bees. Apart from Encarsia the highest value among Hymenoptera is for Bombus terrestris, which many authors in the past have denied the 'right' to fly. Yet, the average lift coefficient of a particularly heavy specimen was only 1-2. It is also seen that the hovering sphingids (Sphinx and Manduca) fall within what can be achieved without involving new principles. This does not exclude that during take-off or escape reactions non-steady situations may arise and be important, as will be discussed later, but it is hardly within the groups mentioned so far that we are likely to find the really interesting exceptions. There are, however, some important deviations like Pieris napi (and probably P. brassicae) and, of course, the chalcid wasp Encarsia formosa. In both cases CL is much too large and in both types the known flight differs from that of normal hovering. For these reasons, the results are shown in parentheses because the simple analysis is hardly justified. As we shall see, the same applies to Syrphinae and Odonata. The results for the bat Plecotus auritus are also in parentheses but they are presumably not far from the true values in spite of the fact that the wings are somewhat flexed during the upstroke.
13
EXB 59
194
TORKEL WEIS-FOGH
Table 6. Corrected values for specific aerodynamic power P * as compared with the result of simple calculations and with a more complete treatment {Weis-Fogh, 1972), together with ratio w\vg.
P* (cal h"1 per gf weight) ratio 0-22 o-is 0-15
O-II O-I2 O-2O
Amazilia fimbriata Manduca sexta Vetpa crabro Tkeobalduj ortftulotc Aides aegypti Drosophila viriUs
Lift/drag ratio 6:1 6:1 6:1 7:2 7H 7:4
Simple theory 10-9
Corrected
values
24
12
1972 values
22
6-8
93
18
21
15-3 23-S J 4-5
28
19
24
(e) Corrected results: a simple approach In the simplified treatment there are two main variables which have been omitted, the tilt /? of the stroke plane relative to the horizontal and the induced wind w. As we shall now see, the tilt is of relatively small significance whereas the induced wind is a major factor when estimating the aerodynamic bending moment Qa and the power Pa. During true hovering the direction of the induced wind w is vertical, its magnitude does not change significantly with time and, at the level of the beating wings, it equals w* = GI(2npR2). (36) So far, we have assumed in equation (5) that the true relative wind V(t, r) which meets a wing segment is equal and opposite to the instantaneous flapping velocity v(t, r) but, obviously, it is the vector sum of w and v(t, r). In order to arrive at a simple method for estimating the accuracy of the results and to introduce corrections when needed, we must know the ratio between w and v(t, r). In the following we shall use the radius of gyration rg of the second moment S of the wing area as a representative distance, and therefore the flapping speed at that distance vg(t). We then have
Moreover, we shall only consider the most common case of a semi-elliptical wing and only during the middle half of the stroke when 80 % of the lift is being produced. Under these conditions, equation (37) is reduced to w\vg = 2-52 x io x J(G)l(</mR*). (38) Some numerical values are listed in Table 6. It is seen that a ratio of 0-2 is common but in some small insects it may approach 0*1. In similarly built animals the weight G is proportional to the cube of a representative length /, i.e. G oc P, and since (f> tends to be constant and independent of size, the ratio in equation (38) will be size-independent provided that n oc l~i. It will increase in proportion to /i if noc / -1 . As will be shown in another paper, most insects vary the frequency with size as n oc l~\ which tends to give a small w/vg ratio for small insects. In Drosophila, on the other hand, frequency and size vary as n oc l~i. The ratio is therefore similar in hummingbirds and Drosophila. In Table 6 the lift/drag ratio is also given, and the correction procedure is best

Vertical
Horizontal Drag D, simple
Fig. I I . The effect of the induced wind to and of the tilting angle /? on the force system acting on the wing of a Drosophila during the middle part of the stroke and at the distance from the fulcrum corresponding to the radius of gyration r, of the second moment S of the wing area, as compared with the simple system in Fig. i. The L/> ratio ia 1-75, tt>/r, is 0-20 and ft is 20 0 .
illustrated with reference to Fig. 11 for Drosopkila virilis, where wjva = 0-2, L/D = 7:4 and /? = 200. In the simple case L and D are vertical and horizontal respectively, the resulting wind force being a vector from the origin to point A. However, va is actually tilted by 20 and the vertical w must be added to give the resulting relative wind velocity Yg. The true resulting wind force F then has a vertical component which is 7 % larger than L and a drag component in the direction of the wing movement, i.e. in the stroke plane, which is 30 % larger than D, in this case. At the same time Vg is 5 % smaller than vg, and the velocity squared is therefore 10 % smaller than the assumed value. The combined result is that CL has been estimated almost correctly whereas the aerodynamic power Pa has been underestimated by 30%. The exact corrections vary with /?, L\D ratio, and wjvg ratio, but for the material as a whole there is no justification for correcting the CL values, which appear to be valid within 15%. However, the power is always systematically underestimated, and some representative values are compared in Table 6. In this and in the previous table, we use the specific aerodynamic power, or power per unit body weight P j . It is seen that the correction is substantial and that the corrected figures compare reasonably well with the results obtained in a more comprehensive treatment of AmaziUa fimbriata and Drosophila virilis (Weis-Fogh, 1972). It should be noted that the corrections are largest when the LjD ratios are high. It is helpful to consider the more general diagram in Fig. 12, which applies to an LjD ratio of 6:1 and a wjvg ratio of 0-2 for semi-elliptical wings. It is seen that the optimum tilt of the stroke plane with respect to the horizontal is about + 200 and that
13-3
[96
TORKEL WEIS-FOGH
Vertical
-15
Fig. 1 a. General diagram for correcting the average lift coefficient CL and the aerodynamic power Pa when ft varies in animals with semi-elliptical wing contours, an LID ratio of 6:1, and a w/v, ratio of 0-3. The true drag in the stroke plane is seen to be about twice as large as the drag D predicted from the simple theory whereas the lift L is affected to only a small extent.
this does not alter the calculated CL significantly. However, the true drag in the stroke plane is about double that estimated from the simple theory. Consequently, in the following we shall use the corrected values for P%, N and t] in Table 5. They hardly differ by more than 20% from the true values. It should be noted that quite large variations in /3 have relatively little effect and we are therefore justified in disregarding this parameter in a treatment whose aim is a first approximation based on simple expressions. (/) Specific aerodynamic power PJ Of the 30 examples from 28 species in Table 5 the available information was incomplete for the bat Plecotus auritus and the moth Amathes bicolorago, which also had an unusually high lift coefficient (i-6). For the two Pieris butterflies and the chalcid wasp Encarsia formosa we have reasons to believe that the basis assumptions are invalid, so that one cannot include them in the general survey but must discuss them separately. The most surprising result is that the power requirement of the 23 remaining species (24 examples) is very similar and does not seem to vary systematically with size, the

Table 7. Evidence for an elastic system in flying insects
The total mechanical power ouput in the absence of elastic forces P*(B+O is compared with the known metabolic rate (cal h"1 per gf body weight.)
Minimum
197
Dynamic efficiency
(V) P*lttH> 43
Metabolic rate
200 100 300 ICO
mechanical efficiency of muscle

0-22
Physiological feasibility Possible Impossible Possible Impossible Impossible Possible
Amaztiia fimbriata
(hummingbird) Vespa crabro (hornet wasp)
0-51 031 0-30 0-70
58
60 40 56
20
0-58
O-20 0-40 O-49 OI7
Apis melUfica (honey-bee) Aides aegypti (yellow-fever mosquito) Eristalis tenax (drone-fly) Drosopkila virilis (fruit-fly)
o-34
'95
us
120
average being 21 cal h" 1 gf-1. The upper limit is 40 for the heavy lamellicorn beetle Heliocopris and the lower limit is 11 for the large sphingid moth Manduca. This must be accepted as an empirical fact and both the absolute magnitude and the range are consistent with earlier results (Osborne, 1951; Weis-Fogh & Jensen, 1956; Weis-Fogh, 1964). It does not immediately follow from equation (19), but this is because one must also consider the effect of Reynolds number (Re) on the drag coefficient CD, as will be discussed elsewhere (Weis-Fogh, in preparation). It is also consistent with the fact that the specific chemical power consumption, the metabolic rate, is of the same magnitude in small and large flying insects (Weis-Fogh, 1964), and probably in small birds and bats as well. This obviously requires more discussion. If we accept that the small insect Encarsia formosa could produce the necessary lift coefficients shown in parentheses and that the drag coefficients are of equal magnitude, its aerodynamic power requirement would be 10-12 cal h - 1 gf-1 (uncorrected) and therefore be of the same magnitude as in other hovering animals. This came as a surprise.
(g) Dynamic efficiency and elastic forces
Table 5-clearly shows that the ratio N = \Qi\m*xl\Qa\ max is high in almost all insects analysed, with the exception of butterflies and Drosopfala. This means that the dynamic efficiency t] is low and that most species must spend 2-3 times more mechanical energy than needed for flight alone, provided that there is no elastic system present in which the kinetic energy of the wings can be stored and later released. In fact, Table 5 provides strong evidence that an elastic system must be present in insects. If we consider the metabolic rates of the six species for which the data are sufficiently good, Table 7 shows that, as to energetics, flight would not be feasible in Vespa crabro, Aides aegypti and Eristalis tenax unless they could make use of elastic bending moments in the thorax or wings similar to those in the locust Schistocerca gregaria, the moth Sphinx Ugustri and the dragonfly Aeshna grandis (Weis-Fogh, 1961, 1972). As to the honey-bee, Apis meUifica has the highest continuous metabolic rate measured in any insect and we do not known how efficient its wing muscles are; it is a borderline case, while in Drosophila the inertial forces are too small to be significant. The available evidence amounts to a circumstantial proof that as a group, flying insects possess and depend upon elastic forces in order to store and release the kinetic energy of the oscil-
TORKEL WEIS-FOGH B
Fig-* 3 (A) The morphology of the chalcid wasp Encarsiaformosa Gahan, used here for the study of hovering flight in very small insects, (a) Tegula, (a)-(6) length of submarginal vein (d), (c) the marginal vein, and () the tip of the anal vein. On the left-hand side the triangular contour function used in the first estimate of Ci is drawn in broken lines while the rectangular area was used for the second estimate in Table 5 and later. (B) The related but larger species Coccophagus tpectabilii Compere, re-drawn after Compere (1931).
lating wings. Under conditions of normal continuous flight the elastic system resides in the thorax and not in the wings because the wings are only bent elastically and to a significant extent during exceptional circumstances. This does not apply to Encarsia
formosa. (h) Hovering flight of a very small insect
The Problem. The calculated average lift coefficient of about 3 for Encarsiaformosa in Table 5 is much too high to be compatible with steady-state aerodynamics. It may be argued that the flight data in Table 4 are incorrect or misleading. To some extent this may be true because the morphological upstroke is of shorter duration than predicted from an harmonic movement of the wings due to the 'clap' period towards the end of each upstroke (cf. Figs. 14, 15), but the effect could not reduce the lift coefficient to less than about 2. We are therefore faced with the problem that these tiny insects do in fact hover and produce an average lift coefficient of 2-3 at a Reynolds number of between 10 and 20. It should be emphasized that we have not taken into account the brim of hairs and bristles seen in Fig. 13 but only the size and shape of the membranous parts of the wings. At present, we can hardly do better since the hairs are invisible on the available films, and it is doubtful to what extent they play a direct role in the process of lift production, as will be discussed when a new mechanism is proposed in the Discussion (p. 217). The Insect. As to morphology, Fig. 13 shows that Encarsiaformosa Gahan resembles an ordinary four-winged wasp. The most outstanding deviations are shared with other very small insects among Hymenoptera and Coleoptera, in particular the brim of long
199
marginal hairs, the 'stalked' wings, and the lack of any significant wing veins apart from a strong reinforcement of the leading edge (costal veins). In addition, the wing hinge is so constructed that the 'stalked' elastic wing bases can be swung above the dorsum and attain a positional angle of more than 1800 (cf. Fig. 15). The effect is that the two pairs of wings can be ' clapped' closely together above the dorsum in spite of the relatively large distance between the wing fulcra of opposite sides. I have often seen this dorsally ' clapped' posture of the wings in dead or preserved specimens, but it is always present towards the end of the upstroke during active flight, whether the insect moves horizontally forwards or backwards in the air, hovers or is engaged in refined manoeuvres. Some details deserve attention since they may be of functional significance. Fig. 13 was constructed from a set of photomicrographs of specimens embedded in glycerol jelly after mild treatment in lactic acid. Staining with methylene blue revealed that resilin hardly plays any significant role in the wing system so that the main elasticity must reside in the flight muscles and in the hard cuticle of thorax and wings, as in larger Hymenoptera. The inner part of the forewing, from the axillary sclerites and the tegula (a) to point (b) where the marginal vein (c) starts, is not ordinary wing membrane but a rather solid plate of hardened cuticle. I have not been able directly to compare Compere's (1931) description of the forewing in the genus Coccophagus, whose members are somewhat larger (Fig. 13 B), because in Encarsia there is no costal cell and the submarginal vein (d) is the strongly reinforced leading edge itself. It is hollow and carries a row of what appear to be campaniform sensilla at its posterior margin, but the most important characteristics are (1) that it can be twisted about its long axis as a thick-walled tube (pronation and supination) and (2) that it is connected with the posterior part and the short anal vein (e) by means of sclerotized elastic cuticle. The wing base from (a) to (b) is therefore a functional unit which is twisted in basically the same way as the whole wing is in large insects, but the twisting does not necessarily result in a uniform twist of the main wing surface. On the contrary, the marginal vein (c) is not confluent with the submarginal vein (d) and there is a transverse, elastic bending zone from (b) to the tip of the anal vein (e). This means that the outer, main part of the wing surface may tend to be turned as a whole plate (pronated or supinated) when the basal submarginal 'stalk' is twisted. Both the upper and the lower wing surfaces of both pairs of wings are covered with small bristles, about 20 /jm long. They may not have any aerodynamic importance but they would prevent the wings from adhering to other surfaces, including the opposing wing surfaces, during the ' clap' period. Finally, the distance between the wing bases of the stalked hindwings is smaller than that of the forewings. This and the fact that the forewings are moved both up and down and forwards and backwards relative to the body axis require a sliding coupling between the two wing pairs. The coupling consists of the rolled-in hind margin of the forewings (called the retinaculum by Compere, 1931, broken line) and the two upwardly directed hooklets at the tip of the marginal vein of the hindwing. In preserved specimens the hindwings are usually unhooked and held at a different positional angle from that of the forewings, indicating that the hindwings do not merely follow the forewings passively although they move together during flight. On the left-hand wings in Fig. 13 A the triangle drawn in broken lines corresponds to
200
TORKEL WEIS-FOGH
Stroke 2 ends
Stroke 3 starts
16
14
Stroke 4 starts
15
16
17
J
1 Horizontal 2mm Fig. 14. Tracingsfroma slow-motionfilmof a hovering Encarsiaformosa whichfliesslowly backwards and upwards, as described in the text The film speed, 7150 frames/sec, was not sufficiently high to permit unambiguous tracings of the wing contours during the fastest part of the wing stroke, as indicated by the broken lines. The frequency n was 403 see"1.
the contour function used in the first example for Encarsia in Tables 4 and 5. It should take care of the possibility that the marginal hairs may be considered as part of the lift-producing surface. More realistically, the hatched rectangle represents the best simple fit for the case that only the membranous parts matter significantly. It should be noted that only the outer two-thirds of the rectangle is used in the calculations because the wings are 'stalked'. In fact, this refinement makes little difference as to CL but is important for the further treatment of lift production. Thefilms.About 50 slow-motion films were obtained, which assisted in building up a picture of free flight, but only ten gave quantitative information about true hovering.
t was a surprise to find that the movements of Encarsia wings basically resemble those of other insects, small or large. During fast forward flight the body angle is small and the wings beat obliquely up and down. During hovering on the spot, or when flying slowly backwards, Fig. 14 shows that the body is almost vertical and the wings beat almost horizontally. A detailed description is necessary but it must be emphasized that the film speed (7150 frames/see) is insufficient for accurate tracings of the outlines of the wings when they move fastest or are ' shadowed' by the body. In the film no. 47 B a freely flying wasp moved slowly into the field of view from the left to the right and so that it also climbed at an angle of about 450 (Fig. 14). Over four complete strokes the frequency was 403 sec"1 and the translational speed of the body was 9 cm sec"1 in the backwards-upwards direction, or sufficiently small to be ignored in relation to the speed of the wings themselves relative to the air, which exceeds 150 cm sec"1 at the radius of gyration during the downstroke and is significantly higher during the upstroke. However, the fact that the insect climbed not only showed that it could produce lift in excess of what is needed for hovering but it also made it possible to estimate during which part of the stroke lift is produced, by means of measuring the vertical oscillations of the body. The tracings in Fig. 14 are from wingstroke no. 3 and relate to the movements of the wings relative to the body. The vertical and horizontal axes are indicated; and frames are numbered so that no. o is the beginning of the morphological downstroke. The animal is seen at a somewhat oblique angle with its long axis slightly tilted by about 200 towards the reader and so that the left-hand side of the head is above the plane of the paper. The right pair of wings (on the far side of the insect) then appears at or slightly above the level of the head, and the left pair beneath. Sometimes it was not possible to distinguish foreand hind-wings, but whenever the interpretation is in doubt, broken lines are used to outline the wings. The anterior or costal edge of the forewings is drawn in heavy line. The toing stroke. Although forewing and hindwing on each side can separate and sometimes do so, basically they move as a single unit, the pair of hooklets ensuring a sliding anchorage to the retinaculum of the forewing. The stroke in Fig. 14 is covered by 18 frames and starts in no. o with the wings held closely together as two plane plates above the dorsum and with the leading edge at about the same level as the forehead. During the next o* 1 msec the two pairs of wings are 'flungopen' like a book, the hindwings representing the back of the book (nos. 1 and 2). The two pairs then separate and move almost horizontally during the morphological downstroke as if they were ordinary aerofoils working at moderate angles of attack (nos. 3-5), until they pass the midpoint where y = 900. The angle of attack is then increased and the wings continue to move for another 300 but are also swung upwards until the end of the downstroke is reached somewhere between nos. 7 and 8. The morphological upstroke starts partly with a rapid reversal of angular movement which actually leads to an elastic bending of the wings at the transverse bending zone (nos. 9 and 10) and partly with a quick ' flip' gf the main wing surface whereby it is rotated as a whole through a large angle so that the co9tal edge again leads (nos. 7-9). It resembles the flipping of a pancake in the air, hence the term. The rapid morphological upstroke then follows with a tilt y? of the stroke-plane angle of about 300 and elastic straightening of the wings; it ends in a' clap' when the wing tips are well below the level of the head (nos. 13-14). The 'clap' lasts about one quarter to one fifth of the entire cycle and
201
202
TORKEL WEIS-FOGH
Table 8. Vertical movements (incmx io~ 8 ) of a freely hovering, slowly climbing Encarsia formosa, as analysed during three successive complete wing strokes (strokes 1-3), each represented by 17-18 frames (nos. 1-18) and filmed at 7150 framesI'sec
Stroke 1 Downs troke Frames Climb 'Upstroke' Frames Climb 'Clap' Frames Climb Total climb (cm x io-*)
0-7 3
Stroke a
0-7
Stroke 3
o-7 3
8-13
3
8-12 6 13-16
3
16
8-13
3
14-17
0
14-17
0
cannot produce any useful aerodynamic forces, because the two pairs are held closely together, their flat surfaces are vertical, and they are moved vertically upwards until the tips reach the level of the head again (no. 17). Detailed analysis of the movements of the body relative to a co-ordinate system fixed in space showed that in three successive wing strokes in this sequence (strokes 1-3), the vertical movements followed the same pattern, as listed in Table 8. The horizontal movements of the body are of less significance since the speed in this direction is small, the wind resistance is therefore small, and the kinetic energy would tend to blur any variations. The division between ' upstroke' and ' downstroke' cannot be made very accurately, but comparison with Fig. 14 should clarify doubts as to the interpretation. It is seen that both the horizontal downstroke and the tilted upstroke produce lift in excess of the body weight in all three strokes, and in approximately equal amounts. Also, the 'clap' period does not appear to contribute to the vertical force; the apparent exception in stroke 2 is probably caused by the' up shoot' following the exceptionally strong lift during the preceding two phases. In any case the resolution of the photographs is too small to permit any further conclusions about the ' clap' period. However, another important observation is that a substantial amount of climb and therefore of lift is produced between frames o and 3, i.e. well before the wings reach maximum angular velocity during the ' downstroke' and immediately following the 'flinging open' phase. Similarly, it looks as if hit builds up immediately after the 'flip', which initiates the 'upstroke', but the rapid angular movements and the elastic deformations of the wings during this phase prevent further analysis at present. The wing-stroke cycle and the angular movements of the long axis of the forewing are summarized in Fig. 15, which derives from another specimen in film 47A. The insect was photographed from in front and almost along the long axis of the body, the wing-stroke frequency being 374 sec"1. During the period of' clap' the positional angle y is larger than 180, as already explained, and the 'downstroke' follows a course not far from that of an harmonic movement at the overall frequency of oscillation. On the other hand the 'upstroke' is considerably faster since the half-cycle includes both the upwards movement and the ' clap' period. Conclusion. Both these and other films were used to estimate the rate of rotation of the wing plane during the periods of 'flinging open' and of 'flip', but these figures arg

Time: 00
203
60 (ms)
20
40
T3
2
a a a a o
\3
00
12 16 0 4 Frame number 12 16 0
10
Fig. 15. The angular movements of the long axis of the wing relative to the body in Encartia formota with an indication of the clap, the fling and the flip phases. The frequency was lower than in Fig. 14, 374 sec"1, and the film speed 7300 frames/sec.
more relevant when we test the new proposed mechanism for lift production in the Discussion to which these observations lead (p. 216). What should be kept in mind at this stage is that the wings are moved essentially as if ordinary aerodynamic principles apply; the useful force is produced almost at right angles to the movement, in contrast to a principle based upon pushing or rowing, which have often been invoked to explain the flight of very small insects. Our problem can then be re-defined as follows: how is an apparent conventional aerodynamic cross-force, i.e. aerodynamic lift, produced by means of flapping wings under conditions which prevent the building up of sufficient circulation F in the usual way?
(1) Hovering flight of syrphid flies and dragonflies
The flight. The true hover-flies belong to the subfamily Syrphinae. Apart from the frequency quoted by Magnan (1934) for Scaeva pyrastri (190 sec"1) and by Rohdendorf (1958/59, 131-170 sec"1) for various Syrphus species, next to nothing is known about their flight. They are of course easily observed; but, in contrast to the distinct hum produced by flying species of the subfamily Eristalinae (Eristalis, HelophUus, Myiatropa, etc.) and Volucellinae {Vohicella), whose wing-stroke frequencies have often been recorded, the flight sounds emitted by the Syrphinae are very weak. It is also characteristic that they retain an almost horizontal body axis whether they hover or dash off in fast forward flight when disturbed, as mentioned earlier (p. 181). Both in Syrphinae and in Odonata this is probably an adaptation to an adult life entirely
TORKEL WEIS-FOGH
Time: 10 ms between dots Fig. 16. Sound tracings from some of the freely hovering syrphid flies recorded in nature and used in Table 9. (a)-(c) Syrphus balteatus hovering in still air at wing-stroke frequencies 177, 123 and 100 sec" 1 respectively. (<f) Syrphus ribetti at 208 sec"1, (e) Sphaerophoria scripta at 309 gee"1 and (/) at 315 sec"1, (g) Syrphus corollae at 163 sec"1, (h) EristaUs arbustorum at 217 sec"1, and (t) EristaUs tenax at 162 sec"1. Distance between dots on the time calibration axis represents 10 msec.
dependent upon flight for feeding and sexual display. In both groups no time is wasted by adjusting the body angle or the visual field when instant manoeuvres are called for. I have seen a male Sphaerophoria scripta hover motionless in front of a female with the body axis tilted head downwards. During typical hovering in still air the stroke angle, <f>, is surprisingly small, estimated to be about 6o and not exceeding 75 in any case, but no accurate measurements exist. This means that the stroke-arc/wing-chord ratio, A, in equation (35) is only about 1*4 in Syrphinae. The wings appear to beat up and down with a tilt of about 45 to the horizontal and the tips definitely do not follow the usual horizontal figure-of-eight; but, again, future slow-motion films or stroboscopic observations of freely flying insects are needed to elucidate the details. In contrast, my unaided observations on foraging EristaUs, Helophilus and VohtceUa indicate that representatives of these subfamilies make use of normal hovering. In fact, Eristalis can be quite difficult to distinguish by sight or sound from a foraging bee. Wing-sound tracings. The sounds produced by hovering Syrphinae are barely audible to the human ear and they are complex, as is seen from Fig. 16. The upper three recordings (a-c) are from the expert hoverer Syrphus balteatus, which can vary its frequency appreciably. It is 177 see"1 in (a), 123 sec"1 in (b) and only 100 sec -1 in (c). The distance between dots on the time-calibration strip represents 10 msec. At all
Table g. The smallest possible
lift coe#amt
& in howring SyrpWpsRF (Syrphidae)
The eight species of w e hover-flies, Syrphinae, use a stroke angle of 75' or l-. The two specia, of drone free, Eriscalinac, use a larger angle of about 120". The d u r n in p a r e n t h m relate to stroke angles not normally seen during free hovering in nature. Even under the m a s t favourable conditions, steady-state aerody&cs ie insufEci-mt to erpl& the hovering of Syrphw and other Syrphinae.)
Chord,
C
(4
0.3 I 0.34 0.38 0.29 0.34 0.36 0.35 0.37 0.38 0.37 0.36 0.36 0.33 0.36 0.40 040 Average 3'0 0.28 0.26 0.26 0.28 Average 3.1 0.37
0.44
Syrphinae SvrPhvJribsfii
Average 0.66
i?rirtalir mburtoMn B. tmax
1.71 1-38
206
TORKEL WEIS-FOGH
frequencies the pattern is very complex, and this applies to all the recordings including those of Syrphus ribesii (d), S. coroUae (g) and Sphaerophoria scripta (e, / ) . The latter clearly demonstrates that the true wing-stroke frequency of 315 sec"1 in (/) would be estimated to be twice as high if a purely auditory method was used, the so-called tenor error discussed by Sotavalta (1947). No attempt is made in this study to interpret the sound tracings. They are merely used to determine the frequency of the beating wings during true hovering of each individual included in Table 9, but they may offer important clues about the flight mechanism at a later stage. The aerodynamic problem. If an animal were restricted to the use of conventional aerofoil action for hovering, the best possible solution would be (a) to make equal use of upstroke and downstroke and (b) to beat the wings almost horizontally, as during normal hovering. Any substantial deviation would require higher lift coefficients. If therefore we calculate the average CL for the syrphidflieson this assumption, known to be untrue for the Syrphinae, we would arrive at the minimum figures seen in Table 9. The contour function of the wings was a semi-ellipse, and c was estimated from the best fit for each individual insect. The figures in parentheses are the results when the stroke angles <p differ appreciably from the true values. It is seen that in all Syrphinae of the three morphological types CL should be as large as 2-3, i.e. incompatible with steadystate aerodynamics by a factor of at least 2 or 3. In fact the real discrepancy is larger partly because A is so small (1-4) and partly because the wings do not beat in a horizontal plane. Clearly, the non-steady flow patterns present at each end of the stroke in any hovering animal (Weis-Fogh, 1972) must be utilized and must have become dominant in Syrphinae, rather than the steady-state phases which play a major role in most other groups. This extreme situation is probably correlated with the relatively low figures for the wing loading of Syrphinae, 0-7 kgf m~2, which contrasts with those for Eristalinae and Volucellinae, where the load figure is 1-2 kgf m~2 (Magnan, 1934; Rohdendorf, 1958/9). According to Table 9 the larger and relatively more heavy Eristalinae seem to rely mainly on conventional principles. However, within flying insects, and in particular within Diptera, there must clearly be a whole spectrum ranging from the ordinary to the exceptional mode of lift production. As to Odonata, calculations similar to those in Table 9 for Aeshna juncea (based on Sotavalta, 1947) and for Aeshna grandis (Weis-Fogh, 1967) show that hovering dragonflies must rely on non-steady aerodynamics because in both species each of the four wings should operate with a minimum CL of 2-3. Again, the wing-load figure for Odonata is low, o-i-o-6 kgf m~2 (Mullenhoff, 1885). In hovering Syrphinae and Odonata there is no 'flinging open' phase, but at each end of the wing-stroke there must be a rapid rotation of the wing surface about its long axis, or a phase corresponding in principle to the 'flip' of the main part of the wing surface in the chalcid wasp.
DISCUSSION
This investigation represents a first-order approximation so that refinements and supposedly second-order effects have been avoided deliberately in order to make the survey manageable in terms of labour and also to discover new principles and solutions
207
Imich may have been overlooked in previous studies. In fact, the conclusions arrived at here and in the previous sections should make it clear that we are now beginning to understand the major mechanisms in hovering flight in terms not only of conventional aerodynamics and mechanics but also of new and maybe hitherto unknown principles. In any given animal species the net result may depend on a complicated combination of mechanisms, as is so often the case in biological systems where countless generations have been selected for fitness in the course of evolution. Since late Devonian, about 350 million years ago, nature has explored aerial locomotion in all its aspects and over a great range of size and form. Of the 1 million known animal species, living and fossil, 750000 are winged insects.
(a) Fast forward flight
During recent years there has been rapid progress in our understanding of the gliding and soaring flight of birds (Pennycuick, 1968 b, 1972 a; Tucker & Parrot, 1970), bats (Pennycuick, 1971), of the extinct pterosaurs (Bramwell, 1971) and of some insects (Jensen, 1956; Nachtigall, 1967; R. A. Norberg, 1972). Similarly, powered flight has been studied in detail in birds (Pennycuick, 1968a, 1969; Tucker, 1968; Bilo, 1971, 1972; Nachtigall & Kempf, 1971), in bats (Norberg, 1970; Thomas & Suthers, 1972) and insects (Weis-Fogh & Jensen, 1956; Jensen, 1956; Weis-Fogh, 1956; Nachtigall, 1966; Wood, 1970, 1972). The list is far from complete, but three main conclusions are either borne out or inherent in most results and arguments presented so far: (1) that the basic mechanism is similar to that first proposed by Lilienthal (1889) for large birds and later found in locusts (Jensen, 1956) as discussed elsewhere (Weis-Fogh, 1961), (2) that, as to quantities, fast forward flight in animals is based mainly upon steadystate aerodynamics (Brown, 1948, 1951, 1953; Weis-Fogh & Jensen, 1956; Jensen, 1956; Pennycuick, 1968a, b), and (3) that many refinements are present within the framework stated under (1) and (2) which can be considered to be consequences of refined kinematics seen in birds, bats and insects (cf. Nachtigall, 1966; Nachtigall & Kempf, 1971; Ruppel, 1971; Bilo, 1972). In a recent book Pennycuick (1972ft) has provided a synthesis of several aspects of vertebrate flight. Although some of his points are arguable, the general conclusions and line of thoughts provide an admirably clear picture of the present state. In contrast to this there have been claims in the literature that some insects with high wing loadings, in particular lamellicorn beetles, must make use of non-steady principles to a major extent (Osborne, 1951 ;Bennett, 1966,1970). We have seen that, although no new principles have actually been proposed but must be related to a high rate of wing twist in some way (Bennett, 1970), the relatively heavy beetles and bees need not necessarily use non-steady aerodynamics, not even when they hover. One of the results of the present study is that exceptions to this rule (i.e. steady-state aerodynamics) are to be found among the insects (and birds?) with a low rather than a high wing loading, as is clearly demonstrated in Table 10. In spite of considerable progress in recent years it still remains true that the flying desert locust Schistocerca gregaria is the best understood and analysed example of fast forward flight, and we should therefore briefly consider Martin Jensen's results from 1956 on the basis of our analysis of the wing movements seen in Fig. 17 (not previously published). It represents traces of the sequence of 18 equally spaced flash photographs j 1 o~6 sec) from stroboscopic film I. The wing-stroke frequency n was 17-5 sec"1 and all
208
TORKEL WEIS-FOGH Table 10. The load figure, i.e. {total weight) I (sustaining wing area), in flying animals
Based upon MUllenhoff (1885), Magnan (1934), Sotavalta (1947), Greenewalt (1962) and present study. () indicates that most of the species make use of normal hovering, () that unusual principles are likely to be important (in the case of DrotophUa to an unknown extent.) Range (kgf/m) Bats Large Up to 8 Medium 2-3 Small* 1-2 Birds Some ducks, Otis, Grus Hens and fowls Vultures and eagles Pigeons and crows Small passerines Hummingbirds* Swifts, swallows, bee-eaters* Insects Coleoptera, Lamellicomia* Hymenoptera, Vespoidea, Apoidea* Diptera, large Brachycera and Cyclorrhapha* Lepidoptera, Sphingidae* Lepidoptera, Noctuidea* Coleoptera, many groups* Diptera, Syrphinae** Odonata** Drotoph&a virilis** Chalcid wasp, Encarsia formosa** Lepidoptera, Rhopalocera" 11-17 8-14 S-io 3-5
2-S 2-3
1-3-2-5 1 2-4-1
0-8-4-s
0-5-2-0 0-4-1-2 0-3-0-6 o-i-o-6
0-3-1-1
o-i-o-6 o-35
O-I2
O-O4-O-2
other flight parameters also appeared to be normal for steady horizontal flight at 3'jm sec"1. The average vertical lifting force was 2-23 gf as measured on the aerodynamic balance, or 97% of the body weight, while the detailed analysis based upon steady-state aerodynamics resulted in 2-17 gf. This discrepancy (3%) is well within the accuracy of measurements and the error arising horn, predictable non-steady effects (Jensen, 1956). The same applies to the measured and calculated average horizontal thrust which is much smaller (o-iogf and 0-14 gf respectively). We can therefore express confidence in the flight situation itself as well as in the method of approach. The numbering of the frames starts with the left forewing at its top position (no. 1) when the hind wing has already started its downstroke. The forewing is marked by means of two very fine white hairs placed normal to the long axis of the wing, one near the tip and the other further towards the fulcrum, where as distinct rear flap is present in the form of the stiff vannal area of the forewings (corresponding to the flexible fan-shaped vannal membrane of the hind wing). The hair was cut at the natural line of bending to permit free operation of the flap. Apart from the downward angular movements of the whole wing the downstroke is characterized by a nose-down twist of the wing, a pronation. This deformation is the result of active movements at the wing base and results in an almost linear twist 8 with distance from the fulcrum. Furthermore, when the forewing approaches the horizontal position and the angular velocity has reached its maximum and begins to decline again, the vannal flap is automatically tilted downwards, altering the wing profile and increasing the lift coefficient (Jensen
209
B Fig. 17. Exact tracings from our original him used Dy Martin Jensen (.1950) tor his analysis of flight no. I in a female Sckistocerca gregaria. The numbering starts when the forewing begins the downstroke and is being pronated, i.e. when the hind wing has already advanced considerably. (A) Downstroke and pronation, (B) upstroke and supination. The upper wing surfaces are dotted and the vannalflapof the forewing is indicated by transverse hatching. The indicators on the wings are minute hairs with white tips and are clearly visible in the films. A typical example of fast forward flight (330 cm s~l) in a large insect (weight 2-30 gf). Note that the elastic bending of the forewing is only slight and can be ignored in the aerodynamic analysis whereas the vannal area of the hindwing is moulded by wind and mass forces.
1956). The upstroke starts with a rapid reversal of the twist, a supination (between nos. 12 and 13), a flattening out of the wing (no. 13), and shortly after an upwards tilt of the flap (nos. 14-16). This gives rise to the so-called Z-profile which offers little lift and drag (Jensen, 1956). These deformations are always present whenever the locust flies actively, but may of course become exaggerated in still air and under other adverse conditions, as seen in Fig. 18. In the latter case the large stroke angle and the air resistance which meets the wing of a struggling tethered animal can result in visible elastic bending, but this is not the case during normal forward flight (cf. Fig. 17(14) and Fig. 18/). In principle, the hind wings are moved in a similar way although theflexiblevannus tends to obscure the basic similarity. In an extremely detailed study of the movements of the forewings of the migratory locust (Locusta migratoria) by means of stereo-photographs taken from above, Zarnack (1972) has recently claimed that Jensen's kinematic analysis is invalid. What he means is presumably that there were second-order effects not taken into account, such as various elastic bendings, and this is undoubtedly so. However, Zarnack failed to demonstrate the much more important flap and the Zprofile, both of considerable aerodynamic importance, and my own conclusion is that his method cannot be as accurate as he himself claims since innumerable direct observations on three species of locusts (Sckistocerca gregaria, Locusta migratoria and Notnadacris septemfasciata), stroboscopic films from the side (Fig. 17) and flash photographs
14
EXB 59
210
TORKEL WEIS-FOGH
Downstroke
Upstroke
Fig. 18. Extreme deformations of the forewing in Schittocerca gregaria when started in still air and flapping at high stroke angle and frequency. Traced from flash photographs. The transverse lines are white markings of cellulose paint. Elastic deformation is now appreciable at the beginning of the upstroke (cf. Fig. ye).
of ScJristocerca gregaria (Fig. 18) never fail to show these features. There are next to no differences in morphology and flight between the three species. The net result of the movements relative to the air is illustrated in Fig. 19 (reproduced from Jensen, 1956, by combining his figs. Ill, 6 and III, 8). The tracings refer to the tip and to the middle section of a forewing as seen on the unfolded elliptical cylinder, whose axis is the horizontally moving wing base indicated by the broken line. It is seen that the geometrical angle of attack of each section is small and carefully adjusted to the path. It should be noted that Jensen's calculations are based on the true three-dimensional movements and that the mutual interference between the wings was taken into account in estimating the aerodynamic angles of attack. The diagram is presumably valid in principle for most types of fast forward flight. It is also reminiscent of a similar diagram in Fig. 20 for the two-winged fly Phormia regina flying forwards at 2*8 m sec"1 and analysed by Nachtigall (1966), the main differences being (1) a much higher rate of wing twist ddjdt of about 6000 rad sec"1 as against maximally 500 rad sec"1 in Schistocerca, (2) high geometrical angles of attack towards the extremes of each half-stroke where the high twist rates also occur, and (3) a larger ratio of flapping speed to forward speed. These points are all relevant in the discussion of hovering flight but at this point it is obvious that the flight of these two very different insects is based essentially on the same type of movement, and that one can envisage a continuous evolution from the one to the other. This also means that the same individual can utilize basically identical mechanisms for fast and slow forward flight as well as for true hovering.
211
18
13
18
Fig. 19. Developed picture (unfolded elliptical cylinder) of the movements of the tip (A) and of the middle part (B) of the forewing relative to the air in a flying Schutocerca gregaria (reproduced from Jensen, 1956, flight no. I).
(b) 'Delayed' elasticity
The high rate of wing twist may mean that an elastic deformation is propagated at a speed which is relatively small compared with that of the rate of deformation from wing base to tip. In the absence of external forces, the velocity of propagation u of a torsional deformation is 2 = Glp, (39) 3 where p is the density of the material and is about 1 g cm" , while G is the shear modulus. The relationship between G and the Young's modulus E is G = /[2(i +/)], (40) where /i is the Poisson ratio and can be taken to be close to 0-4 in a material like insect cuticle. According to Jensen & Weis Fogh (1962) E for the solid cuticle of locust forewings is 870 kgf mm"2 so that G is 310 kgf mm"1 (about 3100 N mm"1), The maximum speed of propagation in a wing is then u = 5-1 x io 3 cm sec"1. The actual deformation will reach the wing section later than indicated by this figure if the move14-3
212
TORKEL WEIS-FOGH
Fig. 20. A picture similar to that in Fig. 19 but from a forward-flying large Dipteran fly Phorrma regata (redrawn from Nachtigall, 1966, Fig. 36). The upper part of the leading edge is indicated by a triangle.
ment is damped by air or other loads and if the elastic modulus is smaller, as it must be in the soft and flexible parts of the wing membrane. How does this influence flight? In the case of a locust with wing length 5 cm and wing-stroke frequency 20 sec"1, each stroke lasts 50 x io" 3 sec and the propagation time is only 2% of the period. The effect will be insignificant. In the fly Phormia regina the duration of one stroke is 8 x io" 3 sec, of which the twist phases at top and bottom last about 0-5 x io" 3 sec each. With a wing length of 1 cm the propagation time is at least o-2 x io" 3 sec. The same applies to syrphid flies, and it means that if the wing twisting itself is instrumental in creating the aerodynamic flow upon which lift depends in these insects, i.e. if the flip is of importance, the action must be propagated from base to tip with a delay which is appreciable or even of the same magnitude as the duration of the pronation or supination. This may have an important and beneficial consequence for hovering flight in Syrphinae and other insects, as will be discussed later. In the case of the small Chalcid wasp Encarsia formosa, the marginal veins are 0-035 cm long and the total wing length is 0-062 cm. The minimum propagation times are then 0-7 x io~5 and 1*2 x io" 6 sec respectively. During the 'flinging open' phase, the two wings open by at least 6 in io" 5 sec and probably by io on the average (see later). We therefore cannot neglect the propagation time of the torsional movement when dealing with the rapid changes at either end of the wing stroke, but it is unlikely to be of importance during the middle part. It should be noted that in geometrically similar animals the relative importance of the propagation time is independent of size {t oc /) when n oc /-1, as is usually but not

Clap
2I3 End of fling
Fling
<J'
'-.AT &*
r=0
Fig. 31. The fling mechanism in the chalcid wasp Encartia formosa. (A) The start, middle and end of the fling in diagrammatic form, to be compared with Figs. 13-15. (B) The 'flinging open' seen in the direction parallel to the long axes of the wings, during the clap (1), in the process (2) and towards the end with the resulting circulations (3) and shed vortices (4). (C) The beginning of the horizontal, morphological downstroke after the separation of the two pairs of wings and the shedding of tip vortices. (D) The geometry of the opening wings during the fling, as referred to in the text. (E) The propagation of the cleft between the two wing pairs caused by ' delayed' elasticity. The air movements are indicated by thin lines.
214
TORKEL WEIS-FOGH
always the case. The significance of this factor is therefore potentially the same i | small and large flapping animals. (c) A novel mechanism for lift production in Encarsia: the fling The idea. The low Reynolds number makes high lift coefficients impossible and yet CL was found to be about 3. It should also be recalled that lift in excess of the body weight is produced early after the 'flinging open' phase seen in Figs. 14 and 15, hereafter called the fling, namely between frames 3 and 4 or considerably earlier than the attainment of corresponding maximum angular velocity by the horizontally beating wings. This is not consistent with ordinary aerofoil theory and practice. Could it be that the fling is a mechanism for creating circulation round the wings before the right and left pair separate and start to move in a horizontal plane during the morphological downstroke in Fig 21C? At first sight this seems improbable because what is needed is the setting up of a circulation in the sense indicated in Fig. 21 A, and the twisting movements (pronation) are of the opposite sense. We are therefore not dealing with an ordinary Magnus effect. However, the possibility exists that a transient circulation of the desired sense could be induced. Once established it would immediately produce lift normal to the direction of the horizontal translational velocity vt and in an upwards direction. It could then be reinforced to some extent during the stroke, but there would always be a tendency for it to approach the low value corresponding to the small steady-state lift coefficients at the relevant Reynolds number. The mechanism then depends on two opposed wing surfaces placed in air of zero velocity and vorticity. They suddenly split apart along the leading edges by an angular movement around the rear edges which can be represented as a 'hinge' (Fig. 21 A) so that air must flow into the space created between the wings from in front. The opposite flow is prevented by the adjoining rear edges forming the 'hinge'. Immediately after this fling, which corresponds to the pronation in normal flight, the wings on either side separate, each carrying a bound vortex with it. The question is, will this movement result in a circulation of the necessary strength T and the correct sense? It is a problem of flow relative to a solid body which suddenly starts to move (impulsive start) in a viscous incompressible fluid; the resulting motion of the fluid must then be irrotational and without circulation. The point is that the body then breaks into two, each of which carries its own circulation of opposite sign but of equal strength. Feasibility. If theflowpattern is consistent with the formation of two bound vortices of opposite sign, circulation can be created without the formation of any starting vortices and therefore without interference or delay due to them, i.e. without any Wagner effect. However, apart from the flow pattern, a number of other conditions must be fulfilled. During the flinging phase where air must flow into the vacated space from above, i.e. in the right direction, Reynolds number must be sufficiently high so that the energy and vorticity are not degraded into heat almost immediately. Also, the boundary layer must be thin relative to the space available. During an impulsive start the thickness S of the laminar boundary in zero at the start and increases with time approximately as $ V(), (41) where v is the kinematic viscosity (Prandtl, 1952). Examination of the films show that the fling lasts about 3 x io~* sec as a maximum..
215
This corresponds to a thickness of 65 fim. Although the hairs (20 jim long) may increase the effective thickness, the boundary layer should not constitute a serious obstacle. As to Reynolds number (Re) we must analyse the system in more detail during the fling itself. For reasons given later (p. 217) we disregard inflow from the tips and consider only the air streaming in from the area delineated by the two leading edges and the arcs at either end, the leading frame (two-dimensional flow in Fig. 21D). The rate at which the volume of instreaming air changes with time when passing the leading frame is dV/dt and the velocity at the leading frame is uB and at distance r, from the edge it is ,. We have dVldt co) rco , . = ; = (42 ^ Yd> "< = 2 0 Since we may consider ddjdt = was constant and 6 = tot, we have ur = r\tt. (43) The velocity of the entering air is then zero at the trailing edge, where r = o and, more important, inversely proportional to time t. A representative distance r for a triangular section would be r = 2^/3, so that
(Re) = & =
(44)
The major part of the fling occurs in 1-4 x io~* sec and the chord is c = 2-2 x io~a cm. After io~B sec, 7 x io~5 sec and 1-4 x io~* sec (Re) is then 75, 10 and 5 respectively. In other words, at the start of the fling the velocity and (Re) are high, energy can be pumped into the flow system at a high rate by the action of the wings, and the flow is not dominated by viscous forces. It should therefore be possible to calculate the circulation according to the principles for vortex-free flow. Circulation. My first and somewhat naive approach was to estimate the strength F of the circulation by calculating the amount of kinetic energy Ek which the fling could achieve by accelerating the air from zero velocity until the widening space between the wings was filled and the wings separated. This could be taken as a minimum figure which was then related to the kinetic energy Ee of the air in a bound vortex of strength F. Since the lift L per unit length of the wing span is related to the translational flapping velocity vt as (45) L = pvtT, vt could be estimated for the given wing length R, or more accurately for 2/3.R, choosing the flapping velocity at the middle of the rectangular area in Fig. 13. The results were sufliciently encouraging in so far as the calculated velocity vt needed to lift the insect was 260 cm sec"1 while the observed maximum value for the wings is about 180 cm s"1. This means that even according to a crude estimate F is of the right order of magnitude and could more than double the lift coefficient; but it was clearly not sufficient, as could be expected from such a crude approach. However, the analysis also showed that the kinetic energy Ek of the inflowing air Ek oc tojt. (46) This means that the time sequence is important. If the fling is faster, at the start than indicated by the average values used, the performance would be improved. This is true also in a more sophisiticated approach.
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TORKEL WEIS-FOGH
At this stage I consulted Professor Sir James Lighthill, who most kindly discussed these ideas with me and wrote (10 December 1972): I have made calculation of the circulation at the end of the fling phase on the following basis: [cf. Fig. 21B]. The angular velocity (radians per second) with which the two wings ap and aq are opening about the point a is taken as w. Thus I first calculate the circulation around the contours shown [B3], neglecting shedding of vorticity during the fling. This is a classical calculation for vortex-free ('irrotational') flow. The answer is exact: for the geometry shown the circulation is 0-69 we* round both the pencilled contours. If for the moment this value be acepted, then we may consider what happens when a small gap appears between the two wings. This should not make any significant difference to the flow because that flow has zero velocity at the point a on both sides. The circulation round the pencilled contours [thin lines in Fig. 21B] should be unchanged, therefore; and should remain unchanged as they move apart by the usual Helmholtz-Kelvin arguments. I have considered the effect of any vorticity shed at the points p and q during the 'fling' phase. At first sight the shedding of vorticity in the sense indicated [see Fig. 21, B 4] is going to diminish the circulation around the contours shown in the previous figure [B3]. Since the calculation has to be made while the wings are still connected at a, however, those circulations are not around truly closed circuits and so we cannot say for certain whether they are diminished by the exact amount of the circulation around the shed vorticity. A calculation needs, in fact to be made; and this (which I have done) shows, interestingly enough, that /the vorticity shed has passed a considerable distance away from the edge by the end of the 'fling' phase then the circulations around the contours shown in the first figure [B3] are diminished by only a moderate fraction of the circulation around the shed vorticity... This leads me to believe that considerable circulations around the two wings would remain at the end of the 'fling'. Apart from shed vorticity (to which we shall return later, p. 217) and viscous effects we then have, for the Encarsia wing immediately at the end of the fling period, when separation of the wings occurs, for each wing approximately (47) During the first 2^4 x io~* sec the angle d is 1300, so that 1200 in Fig. 21 (B 3) and in the calculations is a conservative estimate. The average angular velocity To is 8 x io 3 rad sec"1 during this period and (11 x io 3 rad sec"1 during the first 1-4 x io~* sec). For a chord c = 2-2 x io~2 cm this gives T = 2-67 cm2 sec"1. According to equation (45) and the values for R and vt used there, the flapping speed vt at which the total lift equals the body weight is then 93 cm sec"1. This speed is achieved at about frame 3 in Figs. 14 and 15, i.e. at the time when the films show that the body begins to gain height (cf. Table 8). There is therefore hardly any doubt that this novel method of creating circulation is sufficient and essential for making a small insect like Encarsia formosa airborne. Details of Professor Lighthill's analysis and calculations, which include some considerations of three dimensional aspects of the fluid flows, will be published elsewhere (Lighthill, 1973). Some possible refinements. According to the relationship in equation (46), the principle of which should remain true, the initial period of the fling is crucial as far as energy transfer is concerned and also with respect to Reynolds number (equation 44). We have already seen that the minimum time needed for a torsion of the anterior submarginal and marginal veins (cf. Fig. 13) to travel from the wing base to the distal end

5
217
0-7 x io~ sec; and if the wing surface is made of a similar material it will take 1-2 x io~ sec to reach the wing tip. During the initial io~* sec the average value of 0) is 11 x io 3 rad sec"1 so that io~* sec corresponds to an angular opening of 6. During the initial and energetically most important phase there is therefore no doubt that the cleft between the wings must open gradually from base to tip, as illustrated in Fig. 21E. The implication is that there is no chance of shedding tip vortices until the cleft has reached the tip. In this context it may well be that the marginal hairs have a specific function - that of preventing or delaying tip-vortex formation. It is instructive to note the difference between the hair brim of Encarsia and that of the closely related but larger Coccophagus (Fig. 13 B). Also, Horridge (1956) has pointed out that the smaller the insect the larger and more prominent the hair brim until the wing of the smallest known winged insects (R = 0-02-0-04 cm) consists of a rod with a brim of hairs. We do not know how such wings are moved or whether these insects can fly as freely as Encarsia (R = 0-06 cm), but one should remember that if circulation can be created by means different from those of ordinary aerofoils, the actual shape of the wing is of little importance. What is needed is an elongated solid body which carries a bound vortex and which is moved relative to the stationary air. The hind wings and the prominent hairs at their rear edge in Encarsia may improve the mechanism because it should be possible to fling open the forewings not as one opens a book, but by pulling the two parallel wing surfaces apart while the hind wings seal the rear entrance. This could be the case during the initial phase and should also make it possible for the animal to exert more control than is otherwise possible. In addition, the long hairs on the hindwings could counteract the unavoidable inflow of air at the proximal wing margin. Other flying animals. It was noted that DrosophUa virilis appears to have a clap and a fling period and for that reason it is marked by a triple asterisk (***) in Tables 4 and 5. Although it is just possible to analyse and understand the flight of Drosophila on the basis of steady-state principles (Weis-Fogh, 1972), the usual mechanism does not leave any safety margin, and it appears highly likely that this and other small insects make use of the fling principle, thereby gaining extra lift and manoeuvrability. The cost involved is unknown at present. As to butterflies (Lepidoptera, Rhopalocera) they take off with the wings often held in the clap position at the start, as already demonstrated in Vanessa io by Magnan (1934, p. XIV). The fact that the lift coefficient of Pieris napi in Table 5 is much too high (2-2) for wings of very low aspect ratio strongly indicates that this group of insects regularly utilize the fling method. As to birds, it is easy to hear one or two distinct claps when a pigeon suddenly takes off from a perch when disturbed. Dr R. H. J. Brown has kindly shown me unpublished slow-motion films of flying pigeons from which it appears that at very low air speeds the wings touch each other at the top, when the sound is produced. They are then flung open in a way similar to the fling in Encarsia. It would be interesting to study this phenomenon in large insects also, and in other birds where it might assist flight particularly during take-off in emergencies. Could it explain some of Dr Wilkin's results referred to on page 182 above? Dimensional considerations. The lift of a wing is L oc G oc P oc vtn, where / is a
218
TORKEL WEIS-FOGH
representative length and vt oc nl<j>. In the case of a fling mechanism T oc /2, wherl w a n . We then have L oc P oc n8/4^, or for the specific lift L* = (L/G) oc / oc tfyl. (47) In a series of geometrically similar animals, and if <j> is constant, the usefulness of the fling mechanism is therefore independent of size (L* oc /) when the frequency varies as n oc /~i. This also happens to be the relationship needed for remaining airborne by conventional means, as is seen from equation (9). The fling method is therefore of equal potential use in animals of different sizes. Most animals have to increase n faster than this when they become smaller because the lift coefficient and the lift/drag ratio decrease at low Reynolds number, as has been discussed above (p. 195), and the most common relationship for a given series of animals is n oc l~x. In this case L* oc t~x and small animals will benefit more from the fling than large ones. It is' clearly desirable to confront these deductions with the various modes of flight seen in nature, but they do indicate that the fling mechanism as a regular component of flapping flight should be more favouable in small than in large animals and most beneficial when the wing loading is relatively small. However, in emergencies and other situations of particularly strain the fling mechanism can be used throughout the size range. (d) The flip mechanism in hover flies Obviously the fling mechanism whereby two sets of wings start by forming one structure which is then suddenly deformed before the sets rapidly move apart cannot explain the flight of the true hover-flies, the Syrphinae. However, there are some similarities, and although the following account is much more speculative than the preceding one it is concluded that we are confronted with a special mechanism which I believe to be novel and which I shall call theflipmechanism. Consider an ordinary insect wing which is stretched out from the body. During a period of twist at the extreme positions, be it pronation or supination, the leading edge is suddenly swung actively, nose-down during pronation and nose-up during supination. In many insects the posterior part of the wing is often flexible and does not follow the rapid movement of the front part at all closely. In Encarsia each effective wing surface on each side consists of a large strong forewing and a smaller hind wing. It appears from some of the films as if the hind wing functionally corresponds to the flexible part in a two-winged insect, and I have therefore adopted the descriptive term ' flip' to describe the sudden twist both in Encarsia and in Syrphinae. The wings of Syrphinae (Fig. 22 A, B) are extremely interesting in this context and differ from other Dipteran wings in several respects. (1) The wing itself has a concentration of strong veins going from base to tip in the anterior part while the posterior part is poorly supplied; this is the' aderarme' wing of Rohdendorf (1958/9). (2) There is a special reinforcement known only in the family Syrphidae, the so-called vena spuria (v.s.). It tends to reinforce the anterior almost triangular part and to provide linear reinforcement parallel to which the wing must bend during pronation and supination, separating the anterior, and the posterior parts funtionally. (3) I have observed that the cuticle of the posterior part is extremely thin, soft and pliable in the living insect, particularly at its free border. (4) One could add a functional peculiarity, namely that I have never caught a flying hover-fly which had damaged wings. This
219
Head
(= 0
r=io-
/=2xlO- 4 s
Fig. 22. The flight of hover-flies, Syrphinae. (A) The smallflyPlatychirui pdtatut Mg., showing the large alula (a/.). (B) The wing of Syrpkus balUatus with a large pterostigma (pt.) and the 'false' vein, vena spuria (v.s.). (C) The proposed ./& mechatwm during pronation preceding the downstroke, as resolved in time so that the effect of 'delayed' elasticity becomes apparent; viewed obliquely from above.
contrasts sharply with the common observation that Hymenoptera, Lepidoptera and many other hovering insects are often seen to hover with tattered or worn wings. (5) Finally, the small rear flap at the wing root, the alula (al. in Fig. 22 A), is unusually large and can be bent upwards or downwards at right angles to the remaining wing surface, presumably by means of a pleuro-axillary muscle. It is probably used for control and manoeuvres, particularly in gusty wind, but since it cannot be of importance for lift generation we shall not analyse it further. As a first approximation the wing can be considered as a plate with a stiff anterior part of roughly triangular shape which can be twisted actively from the base and another posterior part which is pliable and is indicated by transverse hatching in Fig. 22 C. The wing length R in a typical hover-fly is 1 cm, and the propagation of a toreional movement from base to tip therefore lasts at least 2 x icr 4 sec. In fact, it is likely to be longer because the wing has an unusually big pterostigmatic area (Fig. 22 B, pt.) recently shown to be more dense and heavy than the remaining part of the wing, at least in Odonata (R. A. Norberg, 1972). At the typical frequency of 150 sec"1, the
220
TORKEL WEIS-FOGH
wing-stroke period is 6-7 x io" 3 sec of which 7% is used for twisting at either enl (Nachtigall, 1966, in Phormia), or 5 x io" 4 sec for pronation and supination respectively. There must therefore be a significant delay between the onset of twisting at base and tip. Also, the soft posterior membrane has a much lower elastic modulus so that the deformation cannot reach this region until after the active twist is completed, i.e. 6-20 x io~* sec after the onset. The air at the hind margin is then at rest during the twisting phases, the margin representing a stagnation line. On this basis I have attempted to illustrate the probable deformations in Fig. 22 C at the onset of pronation (1) when t = o, (2) when the wave has reached the middle part of the wing at t = io-*sec, and (3) when it arrives at the tip at about t = 2 x io~* sec. I have also indicated the likely air movements. At the start the air is at rest. In (2) the middle part is suddenly bent in a fashion reminiscent of the fling in Encarsia but a circulation in the form of a bound vortex cannot be established unless an opposite posterior vortex of opposite sense is created. It will be noted that at this stage, the tip is not moved so that any shedding of vorticity at the tip will be insignificant. The propagation of the twist therefore introduces economy. In (3) the entire wing has undergone twist and has acquired what corresponds to a bound vortex although some useful circulation must now be lost due to the unavoidable tip vortex. When the wing then begins to swing down, leaving the system of posterior vortices behind as if they were ordinary starting vortices, it has already acquired circulation of a strength unrelated to the flapping speed and may produce lift at much lower speeds than envisaged in a steady-state system. It should be mentioned that the vertical induced wind is too small to remove the trailing vortex significantly during the time occupied by the twisting movements. In this mechanism the soft posterior membrane is essential for the initial distribution of the two vortices relative to the wing, the anterior one bound to the wing and the posterior one free. If the posterior part were stiff and the wing a simple torsion plate, the rear part would swing up as the leading part swings down, the system would be symmetrical about the neutral axis and no useful circulation could build up before the translation starts.' Once the real wing is in movement the soft rear flap will also increase the downwash and prevent separation or stall. Without quantitative proof I propose that this is the mechanism which enables Syrphinae and Odonata to obtain the high lift coefficients of 3 or more during hovering flight. The principle could be used by any animal with similar wing characteristics. It differs in principle from the fling mechanism in the way that the posterior vortex cannot be used and tends to counteract the useful bound vortex until the two have become separated in space, and the induced wind is too small (about 50 cm sec"1) to do this during the twisting phases. There is therefore a delayed effect in the aerodynamic system and the two vortices must become separated by translation in order to obtain full lift. The similarity between the fling and the flip is that the transient vortices are produced by the active wing twisting. Future observations and experiments are needed to verify this. At present we have none. Let us now consider the entire wing stroke during hovering and, for the sake of simplicity, assume that pronation (J>) and supination (s) in Fig. 23 produce similar effects at the top and bottom of the stroke. The lift L is always at right angles to the relative wind, the drag D is in the direction of the wind, and the sense of the circula-
221
Dying vortex
Dying vortex
'Normal lift 'Normal' lift
Fig. 23. The effect of the proposed flip mechanism when the wing-tip curve changes in a horizontally hovering syrphine fly (A). The sense of the twist rotation is indicated during pronation (p) and supination (), the resulting bound vortex having the opposite sense (thin lines). In (B) and (C) the wings beat vertically up and down and a strong forward thrust results but no vertical force. In (D) and (E) the fly would experience a steep climb, and in (F) it could hover on the spot, as discussed in the text. (G) is an attempt to combine the non-steady flip mechanism with steady-state phases.
222
TORKEL WEIS-FOGH
tion caused by the flip action is opposite to that of the original wing twist. The flj hovers with horizontal body as in (A). If the wings merely beat up and down in a vertical plane, there will be no net vertical force because the drag components will cancel each other. However, during both downstroke and upstroke there is a strong forwards horizontal thrust caused by the ' flip' lift. If the stroke plane is tilted as in (D) and (E), the lift always points forwards and upwards and the fly will experience a steep climb. True hovering based exclusively on the flip mechanism can be achieved if the wing tip rotates anti-clockwise as in Fig. 23 F and so that the bound vortices die out during the vertical parts of the stroke. In Fig. 23 G I have indicated how the flip mechanism is probably combined with normal aerofoil action in hover-flies and dragonflies during hovering on the spot. It should be noted that during truly non-steady periods the angle of attack is of little importance, and this may well be reflected in NachtingalTa (1966) results from the blow-fly Phormia regina reproduced as Fig. 20, particularly when one examines the large angles towards each end of the stroke. Another point is that even small alterations in wing-tip curve would have quick and drastic effects, as is needed for the supreme control seen in hover-flies. Such changes are well known from Hollick's (1940) studies on the fly Muscma stabulans and are found in most medium-sized Hymenoptera and Diptera. It would be extremely interesting to study free unimpeded flight of hover-flies by means of high-speed cinematography. As far as I am aware, the solution deduced here for the flight of hover-flies (Syrphinae) is new. It does not involve refined adjustments of the angle of attack and requires basically two mechanisms to explain both the act of hovering and the extreme and rapid manoeuvres, namely that the basic flip mechanism dominates the aerodynamic system and that the animal exerts rapid control over the wing-tip path, as is the case in other Diptera (Hollick, 1940). This being fulfilled, the insect can do almost anything without altering the body axis, from hovering to a brisk forward dash, a steep climb or a fall. In a gust of wind the alula could be operated differentially on the two sides so that turning in the yawing plane could be added to the repertoire without interfering with the basic control of flight.
(e) General discussion of hovering flight
At present it is not possible to provide a comprehensive picture but only to indicate the interrelationships which have been established so far, or at least strongly suggested, and to indicate some problems which require a new theoretical or experimental approach. Aerodynamics. A major conclusion from this study is that most insects perform normal hovering on the basis of the well-established principles of steady-state flow, i.e. normal aerofoil action. This implies that the analytical procedure and the mathematical apparatus presented here provide a realistic approach both in qualitative and quantitative terms. However, one must also realize that any type of flapping flight also involves non-steady periods, particularly at the reversal points where active pronation and supination occur. The higher the forward speed and the lower the wing stroke frequency, the smaller is the required rate of wing twisting and the smaller the relative importance of non-steady phenomena, as in a fast flying pigeon, and the empirical observations indicate that when the wing loading exceeds 1 kgf/ma, the steady-state
223
Principles seem to prevail even in hovering animals. The approach illustrated in Fig. 5 and in Table 5 should then provide figures of the correct magnitude, although many modifications can be expected in the light of more detailed knowledge. One of the most urgent requirements is to obtain reliable steady-state lift/drag diagrams of real animal wings, particularly from insects, and measured at the right Reynolds numbers and in a wind field similar to that produced by a truly hovering animal. In addition to the conventional type of hovering we have analysed two novel nonsteady principles for generating lift, the fling and the flip mechanisms, which appear to have become dominant in certain insects with small wing loadings. In both types circulation is set up as a consequence ot the rapid twisting at the reversal pointe in the wing path and before the wing as a whole gains speed relative to the stationary air. The essential difference between the steady and non-steady phases is therefore not that flight depends on an aerodynamic cross-force, or lift, in the first case and on an entirely different mechanism in the other cases; under the conditions considered here, lift depends on the establishment of circulation in the form of a bound vortex round the moving wing. The difference is that a normal aerofoil or wing induces circulation and maintains it as a consequence of viscous shearing forces, particularly in the air passing the rear edge, whenever the wing is moved relative to the surrounding air. Once the circulation has built up and the initial phase is over (Wagner effect) the steady-state implies that the circulation is maintained by the energy dissipated continuously in the shear system. In the case of the non-steady fling and flip mechanisms the vortex patterns which lead to circulations are created prior to and independent of the translation of the wing through the air. These vortex patterns would of course be useless unless they became superimposed upon the subsequent translational movement; but, once established, the air speed of the wing can be reduced relative to what is needed in the steady-state case and for as long as the initial bound vortex remains of significant strength. After it has died out, circulation then has to be maintained by the conventional mechanism. During slow forward flight and hovering the non-steady phases in birds, bats and insects occupy a significant part of the wing-stroke cycle and, as we have seen, they could be put to good use in some cases. Many new examples will undoubtedly come to light in the future. The way in which the aerodynamic bending moment varies with the positional angle in work diagrams like that in Fig. 5 will then be somewhat modified in that the curve ABC will become steeper at the ends and more flat towards the middle, even upwards concave in extreme cases. This will increase the inertial and elastic forces involved in starting a new half-stroke but it does not necessarily result in an increase of the enclosed area, i.e. in a larger aerodynamic power output, because the downwards momentum needed to hover remains the same, in accordance with the experimental results of Wood (1970, 1972). As we have seen, the elastic system of insect wings tends to minimize vortex shedding so that the efficiency may remain reasonably high. If, then, the pronation and supination are used directly to create useful circulation in addition to that obtained by normal aerofoil action, the average coefficient of lift CL as estimated from equation (9) will become reduced. The values in Table 5 are therefore likely to be too large, particularly at low wing loads, because this factor cannot be taken into account until we know more. Drosophila is a good example and has already been discussed (p. 217). What is needed now are new theoretical and
224
TOKKEL W E I S - F O G H
experimental studies on non-steady flow situations as exemplified by the fling and flijl mechanisms, but not necessarily confined to them. It should be recalled that the dimensional analyses show that theflingand flip mechanisms are potentially useful in larger animals also and that they are not in conflict with the conventional mode. It is therefore realistic to proceed with a general dimensional analysis of flapping flight to be presented in another context (paper in preparation). Wing inertia and elastic forces. The same applies to the bending moments caused by the acceleration of the wing mass and by the elastic deformations. So far, the details of the elastic system only seems to have been analysed in the desert locust Schistocerca gregaria (Weis-Fogh, unpublished) and to some extent in Diptera and Hymenoptera (Pringle, 1957, 1968) but it is now clear that similar principles apply to insects in general. However, it is necessary to consider the possible effects of non-steady aerodynamics in this context. rt will be remembered that the inertial torque Q* is relatively insignificant in Drosophila (Weis-Fogh, 1972, and present study). The same applies to the true hover-flies. In typical Syrphinae the wings are relatively very light (0-7% of body weight) and the stroke angles and frequencies are low. When the moment of inertia I was calculated from equation (27) and the ratio iV from equation (23), it was found that the ratio between maximum inertial and aerodynamic bending moments is considerably smaller than unity. This means that in Drosophila and in the Syrphinae the phase lag between tension and shortening essential for the operation of fibrillar muscles (Pringle, 1957, 1967) cannot be provided by wing inertia. At least in the case of Syrphinae it is unlikely that the effective lever arm of the wings changes much with position because of the small stroke angle, and the problem is how a phase lag is introduced. It is possible, however, that a substantial muscle tension is needed to twist the wings against both elastic and aerodynamic forces so that pronation and supination could provide the necessary lag as well as producing useful aerodynamic work. During the evolution of conventional into non-conventional types, or vice versa, there need therefore not have been any real change with respect to myofibrillar mechanism. In Hymenoptera and Diptera the wing twisting is an automatic consequence of the deformation of the elastic thoracic box caused by the strong indirect wing muscles (Pringle, 1957, 1968). In Odonata, Lepidoptera and Coleoptera wing twisting is directly controlled by the basalar and subalar muscles which are well developed in all three orders. We have seen that the anisopterous dragonflies and the Lepidoptera Rhopalocera (butterflies) actually make use of non-steady principles in spite of their relatively large size. The fling phase during the start of flight in Venessa to has already been mentioned, but it may also be significant that Magnan (1934, fig. 174) observed an open wing-tip loop in Aeshna mixta very similar to the one indicated in Fig. 23 G as being optimal for hovering flight based on the flip mechanism. Apart from Hymenoptera the other very smallflyinginsects belong to the Coleoptera and the Thysanoptera and in both orders the basalar muscles are well developed (Matsuda, 1970). There is hardly any doubt that further studies on the dynamics of the flight system in insects will lead to many reinterpretations of the functional morphology of winged insects. The same applies in principle to birds and bats. Apart from propagation of the heart pulse in vertebrate arteries, one does not norm-
225
Illy have to take the speed of propagation of elastic deformations into account when dealing with animal dynamics. However, this factor is important for the non-steady aerodynamics in insects and probably also in the wings of birds and bats. The wing system of insects is often referred to as a mechanical resonant system without further qualifications. This is obviously not true in species with synchronous or non-fibriilar muscles where the rhythm is determined ultimately by the central nervous system, although natural selection would tend to optimize inertial and elastic forces one way or the other. In species with myogenic or fibrillar wing muscles there is no direct nervous control of the fundamental frequency n, and one would expect that n oc J~i provided that the stiffness of the elastic system remains independent of n. At least, this relationship should hold in species with large inertial bending moments such as EristaUs and Calliphora (but not necessarily in Syrphinae). It is therefore disconcerting that Danzer (1956) found that n oc /-- 22 in Calliphora erythrocephala in experiments where the moment of inertia was reduced by shortening the wings. However, he calculated Im from the wrong assumption that the mass was distributed elliptically. For some years Dr Machin in this Department has conducted class experiments with this species and confirmed the predicted relationship, whether Im was found by means of equation (30) or directly by weighing, a minor deviation being caused by the fact that the stiffness of the active flight muscles decreases with increasing frequency (Machin & Pringle, i960). There is therefore good reason to assume that the elastic system is both necessary and efficient in insects whether powered by neurogenic or myogenic muscles.
SUMMARY
1. On the assumption that steady-state aerodynamics applies, simple analytical expressions are derived for the average lift coefficient, Reynolds number, the aerodynamic power, the moment of inertia of the wing mass and the dynamic efficiency in animals which perform normal hovering with horizontally beating wings. 2. The majority of hovering animals, including large lamellicorn beetles and sphingid moths, depend mainly on normal aerofoil action. However, in some groups with wing loading less than 10 N m~2 (1 kgf m~2), non-steady aerodynamics must play a major role, namely in very small insects at low Reynolds number, in true hover-flies (Syrphinae), in large dragonflies (Odonata) and in many butterflies (Lepidoptera Rhopalocera). 3. The specific aerodynamic power ranges between 1-3 and 4 7 W N " 1 (1140 cal h" 1 gf"1) but power output does not vary systematically with size, inter alia because the lift/drag ratio deteriorates at low Reynolds number. 4. Comparisons between metabolic rate, aerodynamic power and dynamic efficiency show that the majority of insects require and depend upon an effective elastic system in the thorax which counteracts the bending moments caused by wing inertia. 5. The free flight of a very small chalcid wasp Encarsiaformosa has been analysed by means of slow-motion films. At this low Reynolds number (10-20), the high lift coefficient of 2 or 3 is not possible with steady-state aerodynamics and the wasp must depend almost entirely on non-steady flow patterns.
15 B I B 59
226
TORKEL WEIS-FOGH
6. The wings of Encarsia are moved almost horizontally during hovering, the bod' being vertical, and there are three unusual phases in the wing stroke: the clap, the fling and the flip. In the clap the wings are brought together at the top of the morphological upstroke. In the fling, which is a pronation at the beginning of the morphological downstroke, the opposed wings are flung open like a book, hinging about their posterior margins. In the flip, which is a supination at the beginning of the morphological upstroke, the wings are rapidly twisted through about 1800. 7. The fling is a hitherto undescribed mechanism for creating lift and for setting up the appropriate circulation over the wing in anticipation of the downstroke. In the case of Encarsia the calculated and observed wing velocities at which lift equals body weight are in agreement, and lift is produced almost instantaneously from the beginning of the downstroke and without any Wagner effect. The fling mechanism seems to be involved in the normal flight of butterflies and possibly of Drosophila and other small insects. Dimensional and other considerations show that it could be a useful mechanism in birds and bats during take-off and in emergencies. 8. The flip is also believed to be a means of setting up an appropriate circulation around the wing, which has hitherto escaped attention; but its operation is less well understood. It is not confined to Encarsia but operates in other insects, not only at the beginning of the upstroke (supination) but also at the beginning of the downstroke where a flip (pronation) replaces the clap and fling of Encarsia. A study of freely flying hover-flies strongly indicates that the Syrphinae (and Odonata) depend almost entirely upon the flip mechanism when hovering. In the case of these insects a transient circulation is presumed to be set up before the translation of the wing through the air, by the rapid pronation (or supination) which affects the stiff anterior margin before the soft posterior portions of the wing. In the flip mechanism vortices of opposite sense must be shed, and a Wagner effect must be present. 9. In some hovering insects the wing twistings occur so rapidly that the speed of propagation of the elastic torsional wave from base to tip plays a significant role and appears to introduce beneficial effects. 10. Non-steady periods, particularly flip effects, are present in all flapping animals and they will modify and become superimposed upon the steady-state pattern as described by the mathematical model presented here. However, the accumulated evidence indicates that the majority of hovering animals conform reasonably well with that model. 11. Many new types of analysis are indicated in the text and are now open for future theoretical and experimental research. This study is dedicated to the memory of my late wife, Hanne Weis-Fogh, n6e Heckscher, who helped, encouraged and inspired me throughout twenty-five years of research, until she died suddenly in a car accident on 17 April 1971. I am greatly indebted to Professor Sir James Lighthill, F.R.S., not only for permission to quote his unpublished caclulations, but also for the open mind with which he received me. I thank Professor J. W. S. Pringle, F.R.S., for information on large beetles, Dr R. H. J. Brown for showing me unpublished slow-motion exposures of flying pigeons, Professor W. D. Biggs for information about wave propagation in solids, Dr Martin Jensen for permission to reproduce Fig. 19, Professor W. NachtigalJ
227
lor permission to reproduce Fig. 20, Dr K. E. Machin for numerous discussions, and Dr J. Smart for expert advice on insect wings. It would have been impossible to provide the slow-motion films and the sound recordings without the enthusiastic assistance of Messrs B. J. Fuller, G. G. Runnalls and D. M. Unwin. I also thank Mr J. W. Rodford for drawing most of the figures and Mr R. T. Hughes for checking the references.
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APPENDIX I
Main symbols and some constants and conversion factors between the technical gram force - centimetre - second system (gf - cm - s) used here and the SI system of measurement.
Superscript 1 m Specific, i.e. per unit weight. Subscript
a e
aerodynamic elastic
D drag
Symbols and constants
g i L
gyration inertial lift
cD c cL
D G I L I M N n
P Q
R r
coefficient of drag coefficient of lift wing chord, as defined aerodynamic drag weight of animal moment of inertia aerodynamic lift representative length mass ratio, |j5<|niax/|Qalmax wing-stroke frequency power bending moment wing length distance, radius
(Re) Reynolds number 5 acR3, second moment of wing areas about fulcra T TCR*, third moment of wing areas about fulcra t time V,v,u velocity (volume) W work w induced wind P angle of tilt F circulation 7 positional angle 7] dynamic efficiency 6 angle of twist A ratio, stroke-arc/wing-chord
230
TORKEL WEIS-FOGH kinematic viscosity: for air at 15 C and 760 mmHg, v = 0-14 cm2 s"1 = 0-14 x io~ 4 m 2 s~ 1 . mass density; for air at 15 C and 760 mmHg, p = 1-25 x io~* gf cm"4 sa = 1*23 x io~3 g cm" = 1 -23 kg m~s. shape factor for second moment of wing area. T shape factor for third moment of wing area. <f> wing-stroke angle; 1 rad = 57*3. (o angular velocity; rad a"1. o"
Conversion factors force 1 gf equals 9-81 x io" 3 N. work 1 gf cm equals 9-81 x io" 6 J. power 1 gf cm s"1 equals
9-8IXIO-W
mass density moment of

inertia
specific power
= 8-44 x io~2 cal h" 1 = 1-3 x io~ 7 Hp (int.). 1 cal h" 1 gf"1 = o-1186 W N" 1 .
1 gf cm"1 sa equals 981 g. 1 gf cm"4 s2 equals -3 981 g cm 1 a 1 gf cm s equals si g cma
APPENDIX 2
Second and third moments of the wing area Second moment S The general expression is 5 = crcR3, where cr is the shape factor. It can be found in most engineering handbooks but is derived here for the case of a semi-elliptical wing in which c is the minor axis and R is the major semi-axis. The semi-chord then varies with the distance r from the fulcrum as (48) Wr) = {cl2R){R*-r*)i. If we calculate the moment about c for two entire wings (or for two whole pairs of wings),
and therefore cr = \ny as in Table 1. Third moment T For any wing T is defined as
r-B = TCR*, J r-0 (So)
where T is the shape factor. In the case of a semi-elliptical wing as before, and if we consider two entire wings,
where ^ is the shape factor. It was calculated in a similar way for the other wing shapes in Table 1.

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7. Exp. Biol. (1973). 59.

169-230 With 23 text-figures Printed in Great Britain

Flight fitness in hovering animals

Flight fitness in hovering animals

c, Factor in eq. (116)

0-690 6-6 0693

Qa{t) = \pCD x I]]**)**

which, when combined with equation (14), gives

Flight fitness in hovering animals

30 40 Distance from fulcrum (mm)

Flight fitness in hovering animals

Since the total mass of the plate is M = \pdcR,

Insect order Odonata Coleoptera Lepidoptera

Flight fitness in hovering animals

0-4 0-6 0-8 Relative distance from tip,

Flight fitness in hovering animals

Flight fitness in hovering animals

Flight fitness in hovering animals

Flight fitness in hovering animals

Flight fitness in hovering animals

Table 4. Flaght data used in this d y &

Bats: Plccotus m ' t u s *

Birds: hummingbirds, Tmhili

AmagiLia jimbriata j-71Niacilir

3'7 5-1 20'0

0.60 0.29 7'8 I 2.8 0.46 0.14

0.04 0.14 1-6 2'12

sphinx ligurtn' Manduca sexto M. swta Mocrogloura stcllatonon Amatha bicololago

Hymenoptera: wasps, been

6I 2 . 7 15.8 13.1 0.85

Bombur t m e r t r i r B. lapidmiur Apir mefijica E w s i a fonnosa (Chalcid wasp)+++ E. f m s a (Chalcid wasp)+++

0.001 0.15 0.056

Bats: P h h u mrthu Birds: hummingbirds, T d i

Metahlic rate during actual flight ( c a l h-l) per gf body weight

11.7 10.9 10.5

11'3 9'4 18.5 20.2 15'7 8.2

Ceionio m a t o Certunbycid species Lepidoptem: butterfliee, moths Finir nap?

(0.4) 3'4 3.6 8.0 5'8

P. brassicae++ sphinx ligurtri Mcmduca sexta

3'7 15.3 23'5 9'3 9'9

1'7 5-6 2.8

Flight fitness in hovering animals

Lift/drag ratio 6:1 6:1 6:1 7:2 7H 7:4

Simple theory 10-9

15-3 23-S J 4-5

Flight fitness in hovering animals

Horizontal Drag D, simple

Flight fitness in hovering animals

mechanical efficiency of muscle

Physiological feasibility Possible Impossible Possible Impossible Impossible Possible

0-51 031 0-30 0-70

Flight fitness in hovering animals

Flight fitness in hovering animals

Flight fitness in hovering animals

Table g. The smallest possible

i?rirtalir mburtoMn B. tmax

Flight fitness in hovering animals

Flight fitness in hovering animals

Flight fitness in hovering animals

(b) 'Delayed' elasticity

Flight fitness in hovering animals

2I3 End of fling

Qa{t) = \pCD x I]])