Wing kinematics and aerodynamic forces in

miniature insect Encarsia formosa in forward

flight
Cite as: Phys. Fluids 33, 021905 (2021); https://doi.org/10.1063/5.0039911
Submitted: 09 December 2020 . Accepted: 10 January 2021 . Published Online: 25 February 2021

Xin Cheng (程欣 ), and Mao Sun (孙茂 )

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Phys. Fluids 33, 021905 (2021); https://doi.org/10.1063/5.0039911 33, 021905

© 2021 Author(s).
 Physics of Fluids ARTICLE scitation.org/journal/phf

Wing kinematics and aerodynamic forces

in miniature insect Encarsia formosa
in forward flight
Cite as: Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911
Submitted: 9 December 2020 . Accepted: 10 January 2021 .
Published Online: 25 February 2021

Xin Cheng (程欣), and Mao Sun (孙茂)a)

AFFILIATIONS
Institute of Fluid Mechanics, Behang University, Beijing 100191, China

a)
Author to whom correspondence should be addressed: m.sun@buaa.edu.cn

ABSTRACT
Miniature insects fly at very low Reynolds numbers, and the effect of air viscosity is large. Previous studies in this area are on hover flight.
Here, we study the forward flight, by measuring the wing kinematics and analyzing the flows of a typical miniature insect (Encarsia formosa,
wing length of about 0.5 mm). In the beginning of the upstroke, the wings quickly accelerate backward at a very large angle of attack and
smash on the air (“impulsive rowing”), generating a large thrust; in the rest of the upstroke, the wings come together and move upward,
slicing through the air and generating a small negative vertical force and negative thrust. In the beginning of the downstroke, the wings fling
open and produce a leading-edge vortex (LEV) on each wing; in the rest of the downstroke, the wings move downward and forward with the
LEV staying attached, generating a large vertical force and some negative thrust. The large thrust produced by the “impulsive rowing” over-
comes the body drag and the negative thrust produced by the wings in the other parts of the flapping cycle; the vertical forces, produced by
the “flinging” and by the downward/forward motion of the wings carrying the LEVs created at the fling, provide the weight supporting force.
That is, the tiny insect overcomes the strong viscous effect by fast smashing the wings on the air, by fast flinging open the wings, and by using
the LEVs created at the fling.
Published under license by AIP Publishing. https://doi.org/10.1063/5.0039911

I. INTRODUCTION aerodynamic forces is the delayed-stall mechanism.10–21 In the

Miniaturization is one of the principal directions of evolution
in insects.1 As a result, many insects are very small. The average
insect-wing length (R) is
5 mm. Near half of the existing winged-
insect species are R between
0.5 mm and
4 mm;2 they are
referred to as miniature insects. The wing length of medium and
large insects is
5 mm–50 mm, e.g., those of fruit-flies, honey bees,
and hawkmoths.2–4
Medium and large insects in normal hovering stroke their wings
back and forth in an approximately horizontal plane [Fig. 1(a)]; the
back and forth strokes are referred to as the upstroke and down-
stroke, respectively.3 It has been shown that in an upstroke and
downstroke, the wings operate at high angle of attack (
35 )5–9
and that a lift and drag are produced, with the drag being a little
smaller than the lift. The lift supports the insect weight, and the
drag in the upstroke and that in the downstroke cancel out each
other. The major aerodynamic mechanism responsible for the
delayed-stall mechanism, a separated leading-edge vortex (LEV)
with large strength attaches to and moves with the wing in the
entire upstroke and downstroke, and the LEV produces a low pres-
sure or suction force on the upper surface of the wing, resulting in
the lift and drag. The mechanism can also be explained using the
vortex dynamic theory: The LEV, together with the root vortex, the
starting vortex, and the tip vortex, forms a vortex loop. As the wing
moves, the attached LEV facilitates a linear growth of the area of
the vortex loop with time, producing a large time rate of change of
the first moment of vorticity (or fluid impulse), hence a large aero-
dynamic force.22–25 A strong LEV attached to the wing is essential
to the delayed-stall mechanism.
The Reynolds number (Re) of the wing in the flapping mode
shown in Fig. 1(a) is defined as
cU 2UfR2 ðr2 =RÞ
Re ¼ ¼ ; (1)
v vðR=cÞ

Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911 33, 021905-1

Published under license by AIP Publishing
 Physics of Fluids
FIG. 1. (a) Wing-tip trajectory (projected onto the symmetrical plane) and the flap-
ping motion of a wing section of a relatively large insect. (b) Those of a tiny wasp,
Encarsia formosa.
where c is the mean chord length of the wing, U is the mean wing
speed at the radius of gyration of the wing (r2), and  is the kinematic
viscosity of the air (here U ¼ 2Ufr2, where U is the stroke amplitude
and f is the wingbeat frequency). Re represents the effect of air viscos-
ity: lower Re means the wing moves in a more viscous flow. As insect
size (R) decreases, U, r2/R, and R/c do not change greatly and f
increases approximately according to R−0.67 (see Ref. 2). Then, Re is
approximately proportional to R1.33. Thus, for small insects (R very
small), Re will be very low, or the viscous effect will be very large. For
the smallest insects such as the tiny wasp Encarsia formosa (wing
length is about 0.5 mm), Re is
10 (see Refs. 3 and 26). At such low
Re, if the flapping mode of the medium and large insects [Fig. 1(a)]
is used, little lift can be generated, while the drag is very large.27,28
This is because with such a large viscous effect, during the forma-
tion of the LEV, the vorticity is highly diffused and the resulting
LEV is very weak.29 Therefore, very small insects must use a “new”
flapping mode.
Recent studies have shown that the flapping mode and force pro-
duction mechanisms of very small insects are very different from those
of their larger counterparts:26,30 the tiny wasp Encarsia formosa and
the thrip Frankliniella occidentalis have a very deep U-shaped upstroke
[Fig. 1(b)], and at the start of the downstroke, the wing performs the
“fling” motion, which was previously discovered by Weis-Fogh.3 The
deep U-shaped upstroke produces large transient drag that points
almost upward, and the “fling” motion, at its later stage, also produces
transient drag pointing almost upward. These two transient drag
FIG. 2. (a) A sketch of the experimental system. (b) Shapes of the model body. (c) Shape of the model wing. (d) Portions of computational grids.
Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911
Published under license by AIP Publishing
ARTICLE scitation.org/journal/phf
forces are produced by fast downward acceleration of the wing, at
nearly 90 angle of attack, similar to the case of a plate rapidly started
at its normal direction.26 These two approximately upward pointing
forces provide the weight-supporting vertical force. The horizontal
components of these two forces have opposite directions and cancel
out each other in a flapping period, and the wingbeat-cycle mean hori-
zontal force is zero, as required for hovering flight.
As far as we know, all the previous studies on the flapping
motion and aerodynamic forces of miniature insects are within the
context of hovering flight.3,26,30–37 In forward flight of miniature
insects, how the flapping-motion pattern and aerodynamic-force
generation would change is unknown. In the present work, we study
the forward flight of a typical miniature insect, the tiny wasp
Encarsia formosa, the hovering flight of which has been studied pre-
viously by many researchers.3,26,34 First, the forward-flight wing
kinematics is measured by using three high-speed cameras
equipped with micro-lenses and extension tubes. Then, the flows
around and aerodynamic forces on the insect are obtained by
numerically solving the Navier–Stokes equations using a well-
tested flow solver. Finally, the flight and force production mecha-
nisms are analyzed, and how the miniature insect overcomes the
large viscous effect is revealed.
II. MODELS AND METHODS
A. Experimental method and insects
The method of measuring the forward-flight wing kinematics is
similar to that used previously by the authors for hovering flight,26
except that the flight chamber in the present study is “longer” than
that used in hover flight: in Ref. 26, the flight chamber is a cubic of the
size of 40  40  40 mm3, while in the present forward-flight study, it
is a cuboid with the size of 80  40  40 mm3 [Fig. 2(a)]. Three
orthogonally aligned synchronized high-speed cameras
(FASTCAM Mini UX100, Photron, Inc., San Diego, CA, USA;
10 000 fps, shutter speed 20 ms, resolution 896  488 pixels)
equipped with 60 mm micro-Nikkon lens and 12 mm extension
tubes are used to film the flight of the insects [Fig. 2(a)].
Quantitative kinematics are extracted from the filmed data by an
interactive graphic user interface developed using MATLAB
(MATLAB V. 7.1, Math Work, Inc., Natrik, MA, USA). The
33, 021905-2
 Physics of Fluids
method has been described in detail in Ref. 26, in which kinematic
data of hovering flight of the same species were measured.
Insects used in the experiment, small wasp Encarsia formosa
Gahan (the same species as that used in the hovering flight study),
were acquired from the Laboratory of Biological Invasion of Institute
of Plant Protection, Chinese Academy of Agricultural Sciences; they
were descendents of wild-caught E. formosa. The wasps were housed
in small chambers in groups of three to five individuals and fed with
20% sugar solution. The experiment was conducted at room tempera-
ture 25  C–27  C and relative humidity 50%–60%.
More details of the experimental setup, the method used to
extract the 3D body and wing kinematics from the filmed data, and
the method used to measure the morphological parameters (wing
length, wing shape, wing area, etc.) can be found in Ref. 26.
B. Flow governing equations, computational model,
and numerical procedure
The governing equations for the flows around the insect are the
incompressible, three-dimensional Navier–Stokes equations. The
dimensionless form of equations is as follows:
r  u ¼ 0; (2)
 
@u 1
þ u  ru ¼ rp þ r2 u; (3)
@s Re
where u is the non-dimensional fluid velocity, p is the non-
dimensional fluid pressure, s is the non-dimensional time, and Re is
the Reynolds number; in the non-dimensionalization, c, U, and c/U
are taken as reference length, velocity, and time, respectively (defini-
tions of c and U have been given in Sec. I).
The body of the insect is modeled as follows: The shape of the
body is obtained from the video images (side and top views of
the body). The cross-section of the body is taken as an ellipse, and the
body model is given by fitting ellipses to the body shape in the video
images. The side top views of the model body are shown in Fig. 2(b).
The wings are modeled as flat plates with rounded leading and trailing
edges, and the thickness of the plate is 0.03c. The planform of the
model wing [Fig. 2(c)] is obtained from the measured wing shape.
There were relative movements between the wings and between
the body and the wings, and therefore the equations were solved over
moving overset grids. In the overset grid system [Fig. 2(d)], there was
a body-fitted curvilinear grid for the body and for each of the left and
the right wings, and there was a background Cartesian grid, which
extends to the far-field boundary of the domain. The background
Cartesian grid was generated algebraically. The wing grid is of O–H
type and that of the body is O–O type, and they were generated by a
Poisson solver. For the wing grids, dynamically deforming grids are
used to treat the time-various bending deformation of the wing near
the end of an upstroke or downstroke. A procedure of combining the
method of the modified trans-finite interpolation and the method of
solving the Poisson equation was used to generate the dynamically
deforming grids; a more detailed description of the procedure can be
found in Du and Sun.38
The CFD solver used to solve the flow equations is an in-house
code developed more than a decade ago13 and has been tested in sev-
eral works of our group,26,33,39,40 using experimental data of various
wing motions (translating at Re ¼ 100;41 rotating at Re ¼ 1000;42
Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911
Published under license by AIP Publishing
ARTICLE scitation.org/journal/phf
revolving with an acceleration phase, a constant-speed phase at Re
¼ 500 and a deceleration phase;43 and flapping44 at Re ¼ 180), and
comparison has also been made with theoretical (Stokes and Oseen
solutions) and measured results for a sphere at very low Re (Re
¼ 5–20). The method has been described in detail in Refs. 26 and 39,
and only an outline of the method is given here. The method is based
on the idea of artificial compressibility, first developed by Rogers
et al.45 for a single grid and then extended by Rogers and Pulliam46 to
overset grids. In the method for a single grid, the time derivatives of
the momentum equations are differenced using a second-order, three-
point backward difference formula. The continuity equation requires
that the divergence of the velocity field is zero. In order to solve the
time discretized momentum equations for a divergence-free velocity at
a new time level, a pseudo-time level is introduced into the momen-
tum equations and a pseudo-time derivative of pressure divided by an
artificial compressibility constant is introduced into the continuity
equation. The resulting system of equations are iterated in pseudo-
time until the pseudo-time derivative of pressure approaches zero;
thus, the divergence of the velocity at the new time level approaches
zero. The derivatives of the viscous fluxes in the momentum equation
are approximated using second-order central differences. For the
derivatives of convective fluxes, upwind differencing based on the
flux-difference splitting technique is used. A third-order upwind
differencing is used at the interior points, and a second-order upwind
differencing is used at points next to boundaries. For the case of overset
grids, the solution method for a single grid is applied to each of the
wing grid and the background grid, and data are interpolated from one
grid to another at the inter-grid boundary points using tri-linear inter-
polation. Details of this algorithm can be found in Refs. 45 and 46.
For the boundary conditions, at the far-field inflow boundary,
the velocity components are specified according to the constant inflow
velocity while pressure is extrapolated from the interior; at the far-
field outflow boundary, pressure is set equal to the inflow pressure
and the velocity is extrapolated from the interior. On the wing sur-
faces, impermeable wall and no-slip boundary conditions are
applied, and the pressure on the boundary is obtained through the
normal component of the momentum equation. The insects move
forward at some constant flight speed, and this value is given at the
inflow boundary.
The grid size used here is the same as that used for hovering flight
in Ref. 26: the wing grid had dimensions 61  91  65 in the normal
direction, around the wing, and in the spanwise direction, respectively
(first layer grid thickness was 0.001c); the body grid had dimensions
99  77  53 along the body, in the azimuthal direction, and in the
normal direction, respectively; the background grid had dimensions
121  121  121 in the x, y, and z directions, respectively. Detailed
grid resolution tests were conducted to ensure that the flow calcula-
tions are grid independent.26
III. RESULTS AND DISCUSSION
A. Wing-flapping motion at forward flight
Constant-speed forward flights of five wasps were captured in
our experiment. These insects are named, respectively, as W1, W2,
W3, W4, and W5. The movies in Fig. 3 (Multimedia view) give the
video sequences for some of the wasps. As an example, the sequences
of the flight of one of the wasps (W1) are shown in Fig. 3.
33, 021905-3
 Physics of Fluids ARTICLE scitation.org/journal/phf

The method for describing the wing motion as a function of time
is the same as that used by Cheng and Sun26 and Lyu et al.30 for minia-
ture insects: For a filmed wasp, data of five flapping cycles were proc-
essed. In each of the five cycles, when the wings are at the end of the
downstroke and at the end of the upstroke, the wing-tip points are
projected onto the plane of symmetry of the insect. A linear regression
line of the projections is then determined. The plane that passes the
wing base and at the same time is parallel to the above line is defined
as the stroke plane. The stroke plane is shown in Fig. 4. In the figure,
(X, Y, Z) is a reference frame with the origin at the wing base: the
X-axis in the horizontal direction and pointing forward, the Z-axis in
the vertical direction and pointing upward, and the Y-axis in the
horizontal direction and pointing sideward (Fig. 4); (x, y, z) is another
reference frame also with the origin at the wing base, the x–y plane
coinciding with and the z-axis normal to the stroke plane, and the
y-axis pointing sideward, coinciding with the Y-axis [Fig. 4(b)]. Both
frames move with the body at constant speed. The stroke plane is
approximately horizontal when the wasp is at hovering flight.26 At
forward flight, it tilted forward by an angle b [see experimental data
in a later part of the present section (Sec. III A)]; b is referred to as
the stroke plane angle [Fig. 4(a)]. The angle between the long-axis
of the body and the horizontal is referred to as the body angle, v
[Fig. 4(a)]. As seen in Fig. 4(a), b + v is the angle between the long-
axis of the body and the stroke plane.
The position (and motion) of a wing relative to the stroke plane
can be determined by three Euler angles and by the maximum bending
displacement of the wing (see Ref. 26 for the detailed description of
wing bending). The Euler angles [Fig. 4(b)] are the positional angle of
wing, / (in the stroke plane), the pitch angle w, and the deviation
angle h; and the maximum bending displacement is denoted by dm.
Figure 5 gives the measured Euler angles and the spanwise bending in
one wingbeat cycle in a forward flying wasp (W1); the corresponding
data of other wasps are given in the Appendix (see Fig. 14). Flight
speed Vf, stroke plane angle b, and body angle v are also obtained
from the measurement. From the measured data, the stroke ampli-
tude Ф (defined as Ф ¼ /max − /min), the wingbeat frequency f,
the advance ratio Vf* (defined as Vf* ¼ Vf/2ФfR), and the Reynolds
number of wing Re (defined in Sec. I) are determined.
Morphological data required for carrying out the flow computation
were measured in the experiment; they are the wing length R, the
area of one wing S, the distance between the two wing roots lr,
and the radius of the second moment of wing area or radius of gyra-
tion r2. These kinematical and morphological data are listed in
Table I.

FIG. 3. Video sequences of W1 at forward flight; three camera views are shown. Times noted are microseconds from the instant when the upstroke starts. Complete video
sequences can be seen in the movies. Movie 1: Flight of W1. The left, middle, and right parts of the movie show the flight captured by the top-view camera and two side-view
cameras, respectively. Playback speed is 15fps,
0.15% of the actual speed of the movie. Movie 2: Flight of W2. The left, middle, and right parts of the movie show the flight
captured by the top-view camera and two side-view cameras, respectively. Playback speed is 15fps,
0.15% of the actual speed of the movie. Movie 3: Flight of W3. The left,
middle, and right parts of the movie show the flight captured by the top-view camera and two side-view cameras, respectively. Playback speed is 15fps,
0.15% of the actual
speed of the movie. Multimedia views: https://doi.org/10.1063/5.0039911.1; https://doi.org/10.1063/5.0039911.2; https://doi.org/10.1063/5.0039911.3

FIG. 4. (a) Reference frames. (b) Euler

angles defining the wing kinematics.

Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911 33, 021905-4

Published under license by AIP Publishing
 Physics of Fluids ARTICLE scitation.org/journal/phf

FIG. 5. (a) Measured Euler angles, /

and h. (b) Measured Euler angle, w, and
maximum bending displacement (dm/R).
Data are for wasp W1 (mean 6 s.d.; n
¼ 5 wingbeats); T, stroke period; R, wing
length.

TABLE I. Flight parameters for the five wasps.

ID Vf (m/s) Vf* f (Hz) Ф (deg) b (deg) v (deg) R (mm) S (mm2) lr (mm) r2 (mm) Re

W1 0.33 0.32 354 140.0 35.9 42 0.59 0.13 0.22 0.38 9.5
W2 0.27 0.25 351 146.8 30.52 45 0.61 0.14 0.23 0.39 10.5
W3 0.32 0.32 349 139.2 33.0 50.5 0.59 0.13 0.22 0.38 9.3
W4 0.25 0.26 328 137.3 22.7 58 0.60 0.13 0.22 0.38 8.9
W5 0.31 0.30 337 136.5 34.3 48 0.64 0.15 0.24 0.41 10.4

Using the data given in Fig. 5 and Table I, the stroke pattern of
the wing motion of W1 is plotted in Fig. 6; those of the other four
wasps are given in the Appendix (see Fig. 15). Figure 6 also shows the
orientation of the wing and the velocity of the wing relative to the air
at various times of the stroke cycle; here, the wing is represented by
the wing section at the radius of gyration of the wing, and the relative
velocity is the vector summation of the velocity due to wing-flapping
(/_ and h)
_ and the velocity due to the insect's forward flight (V_ f ) (see
Fig. 7).
From Fig. 6, it is seen that a flapping cycle can be divided into
four phases. In the first phase (t/T  0–0.25; t denotes time, and T is
the flapping cycle period, T ¼ 1/f), the wings move backward at very
large velocity at large angle of attack (see the left panel of Fig. 6); this
phase is called the “impulsive rowing” phase. In the second phase
(t/T  0.25–0.5), the wings move close to each other at the back of the
insect and at the same time move upward (see the left and right panels
of Fig. 6); this phase is called the closing/up-moving phase. In the third
phase (t/T  0.5–0.65), the wings fast “fling open”3,26 (see the right
panel of Fig. 6). In the fourth or the last phase (t/T  0.65–1), the
wing sweeps downward and forward at large angle of attack (see the
left panel of Fig. 6); this phase is called the downward/forward
sweeping phase. A flapping cycle of insects commonly comprises two
half-strokes, the upstroke and the downstroke. Here, the impulsive
rowing phase and the closing/upward-moving phase are in the
upstroke, and the fling phase and the downward/forward sweeping
phase are in the downstroke.
Comparing the forward flight case in Fig. 6 (left panel) and the
hovering flight case in Fig. 1(b), it is seen that motion of the wing

FIG. 6. Left) Diagram showing the wing-

motion pattern of W1; the dashed curve
indicates the wing-tip trajectory relative to
the body (projected onto the X–Z plane);
black lines indicate the orientation of the
wing at ten time instances, with dots mark-
ing the leading edge; the blue arrows rep-
resent the relative velocity of the wing
(projected on to the X–Z plane). (Right)
Back view of the orientation of the wing at
nine time instances; the blue arrows rep-
resent the relative velocity of the wing
(projected on to the Y–Z plane).

Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911 33, 021905-5

Published under license by AIP Publishing
 Physics of Fluids
FIG. 7. Components of the relative velocity of the wing.
relative to the body is generally similar in the two cases. The major dif-
ference between the two cases is that in the forward flight case, the
body of the insect tilts forward (the body angle is
45 ), while in the
hovering flight case, the body is approximately vertical (the body angle
is
90 ).
B. The aerodynamic forces
The flows around and aerodynamic forces on the insect are com-
puted by solving the incompressible Navier–Stokes equations, as
described in Sec. II. In the present study, the vertical component
(Z-component) of the total aerodynamic force of a wing or the body is
referred to as the vertical force, and the horizontal component
(X-component) is referred to as the horizontal force or thrust; the
Y-component of the force is the side force (note that the side force
of the left wing cancels out that of the right wing). The vertical force
and the horizontal force (thrust) on the two wings are denoted by
Vw and Hw, respectively, and those on the body are denoted by Vb
and Hb, respectively. Figure 8 gives the time histories of the com-
puted aerodynamic forces of the wings and the body in one flapping
cycle for W1; results for the other four individuals are given in the
Appendix (see Fig. 16).
The following is observed from Fig. 8. In the impulsive rowing
phase (t/T  0–0.25), in which the wings move fast backward at high
angle of attack, a very large thrust peak [Fig. 8(b)] and a relatively large
vertical force peak [Fig. 8(a)] are produced. In the closing/upward-
moving phase (t/T  0.25–0.5), in which the wings are close to each
other and move upward, a negative vertical force [Fig. 8(a)] and a neg-
ative horizontal force [Fig. 8(b)] are produced. In the fling phase (t/T
 0.5–0.65), a relatively large positive vertical force [Fig. 8(a)] and
some thrust [Fig. 8(b)] are produced. Finally in the downward/forward
sweeping phase (t/T  0.65–1), a relatively large positive vertical force
[Fig. 8(a)] and negative thrust [Fig. 8(b)] are produced. As for the
body, a small vertical force [Fig. 8(c)] and a relative large negative hori-
zontal force [Fig. 8(d)] are produced.
From the above results, it is seen that the majority of the vertical
force that supports the insect weight is produced by the fling (t/T
 0.5–0.65) and the downward/forward sweeping of the wings (t/T
 0.65–1) (the fling also provides some thrust) and that the majority
Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911
Published under license by AIP Publishing
ARTICLE scitation.org/journal/phf
FIG. 8. The time histories of the computed vertical (a) and horizontal (b) forces of
the wings (c) and the body (d) in one flapping cycle for W1.
of the thrust that propel the insect's forward motion is produced by
the impulsive rowing of the wings (t/T  0–0.25) (the impulsive
rowing also provides some vertical forces). It can also be seen that
in addition to the negative horizontal force on the body (body
drag), in some relatively long periods (t/T  0.25–0.55 and t/T
 0.65–1), the wings also produce a relatively large negative hori-
zontal force (negative thrust). These negative horizontal forces
are mainly overcome by the large thrust in the rowing phase (t/T
 0–0.25). That is, the insect paddles itself through the air like
people rowing a boat.
These can also be seen clearly from the plot of the total aerody-
namic force on the wings (Fig. 9).
C. How the aerodynamic forces are produced
As seen in Sec. III B, large aerodynamic force peaks are produced
during the impulsive rowing phase, the fling phase, and the down-
ward/forward sweeping phase [Figs. 8(a) and 8(b); t/T  0–0.25; t/T
 0.55–1; Fig. 9]. These force peaks provide the weight supporting the
vertical force and the thrust that overcomes the negative horizontal
forces of the wings and the body drag.
Let us define the lift and the drag of the wing as the components
of the total aerodynamic force that are perpendicular and parallel to
the relative velocity of the wing at the radius of gyration of the wing
33, 021905-6
 Physics of Fluids
FIG. 9. The total aerodynamic force on the wings at various times (green arrows)
in one stroke cycle (left panel, side view; right panel, back view).
(r2), respectively. The vertical and horizontal forces of a wing come
from its lift and drag. So, let us look at the lift and drag of the wings
and analyze how they are generated. Figure 10 shows the lift (L) and
drag (D) of the wings of W1 in the flapping cycle.
As seen from Fig. 10, very large drag and much smaller lift are
produced by the wing. That is, the large thrust and large vertical force
in the rowing phase, fling phase, and downward/forward sweeping
phase are mainly contributed by the wing drag. This can be seen more
clearly from Fig. 9: the total aerodynamic force vector is almost in the
opposite direction of the wing velocity vector.
In order to explain how the large wing drag is produced, we
examine the flow field data of the wings. First, we consider the impul-
sive rowing phase (t/T  0–0.25). In this phase, the wings fast acceler-
ate backward at very high angle of attack; i.e., the wings fast smash on
the air. Figure 11 gives the flow field data in this phase. It is seen that
when the wings move, starting from zero velocity, vorticity of opposite
sign is created continuously at the leading and trailing edges and also
FIG. 10. The time histories of the lift L (a) and drag D (b) of the wings of W1 in the
flapping cycle.
Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911
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at the side edges [Fig. 11(a)], similar to that of a plate fast started in its
normal direction. From vortex dynamics theory,47 generation of large
opposite sign vorticity at different locations of the wing will result in
large time rate of change of the first moment of vorticity, giving the
large aerodynamic drag on the wing. The large drag can also be
explained in another simpler way: the wing starts from a relatively
small speed and fast accelerates at very large angle of attack, like a plate
smashes on the fluid, producing a large fluid impulse [Fig. 11(b)]; this
will result in a large positive pressure on its lower surface and a large
suction pressure on its upper surface [Fig. 11(c)], giving in the large
drag.
Next, we look at the fling phase (t/T  0.55–0.65); the flow
field data are plotted in Fig. 12. In the fling phase, the wings fast
rotate about their trailing edges, starting from zero rotational veloc-
ity, also generating large and opposite-sign vorticity at different
locations [Fig. 12(a)]. The wings also smash on the fluid and pro-
duce a large fluid impulse [Fig. 12(b)], giving a large positive pres-
sure on its lower surface and a large suction pressure on its upper
surface [Fig. 12(c)]. The force production mechanism is similar to
that of the impulsive rowing motion. Note that in this phase, a
leading-edge vortex (LEV) is produced by each of the wings [see
the panel in Fig. 12(a) at t/T ¼ 0.64].
Finally, we examine the downward/forward sweeping phase (t/T
 0.65–1); the flow field data are given in Fig. 13. In this phase (which
is subsequent to the fling phase), the LEV produced in the fling phase
attaches to and moves with wing, or the LEV is recaptured by the wing
[Figs. 13(a) and 13(b)]. This is the well-known delayed stall mecha-
nism of aerodynamic force production. That is, the large aerodynamic
force in the downward/forward sweeping phase is due to the LEV or
delayed-stall mechanism. Another way to explain the force production
in this phase is as follows. When the wing having a vortex (the LEV)
on its upper surface, a suction pressure will be produced by the vortex
[see the panels in Fig. 13(c) at t/T ¼ 0.70, 0.75, and 0.80], giving the
wing drag.
As aforementioned, the fling phase and the downward/
forward sweeping phase are the downstroke of the very small wasp
(the fling is at the beginning of the downstroke). For the previously
studied larger insects (no fling at the beginning of a downstroke),
the wing needs to move (azimuthally rotate) about 30 –40 before
the LEV is formed; when the Reynolds number is very low, i.e., the
viscous effect is very large, the vortex formed after such a long
time is rather diffused and cannot be very effective.27–29 Here, to
overcome the large viscous effect, the small wasps “cleverly” use the
fling motion at the beginning of the downstroke. This is a very
interesting point.
We, thus, see that smashing on the air at large acceleration by
the wings is one method used by the tiny wasps to produce a large
aerodynamic force in a very viscous flow: in the impulsive rowing,
very large translational acceleration is used; in the fling, very large
rotational acceleration is used. Another method is using the fast
fling to produce a strong leading-edge vortex and keep the vortex
on the wing in the rest of the downstroke (as aforementioned, in a
normal downstroke used by large insects, when the flow is very
viscous, a strong leading-edge vortex cannot be produced). Using
large acceleration to produce a large aerodynamic force in very vis-
cous flow has also been found in the case of hovering flight of the
tiny wasps in our previous study.26 But at there, the method was
33, 021905-7
 Physics of Fluids
used in vertical-force production. Here, we found that the method
is used in thrust production. Using fast started rotation about
the trailing edge to produce a strong leading-edge vortex in a very
viscous flow and keep the leading-edge vortex in the rest of the
downstroke for large aerodynamic force production is a mechanism
observed for the first time in insect flight.
Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911
Published under license by AIP Publishing
ARTICLE scitation.org/journal/phf
FIG. 11. Vorticity fields, flow fields, and
pressure distribution on the wing surface
in the impulsive rowing phase (t/T
 0–0.25). (a) Isovorticity surface plots of
the wing; vorticity is non-dimensionalized
by U/c, and the magnitude of the non-
dimensional vorticity is 2. (b) Non-
dimensional flow velocity vector plots at
section 0.7R (R is the wing length) at the
corresponding times. (c) Pressure distribu-
tions on the upper surface of the wing
(first row) and the lower surface of the
wing (second row) at the corresponding
times; the pressure is non-
dimensionalized as (p–p1)/0.5qU2, where
p is the pressure, p1 the reference pres-
sure, and q is the air density.
Having explained the aerodynamic-force production mecha-
nisms of the tiny wasps in forward flight, let us consider the aerody-
namic power required to produce the aerodynamic forces. From
the computed aerodynamic force and the wing motion data, the
power used by the wing to generate the forces can be calculated,
and the mean power and mean vertical force (average over one
FIG. 12. Vorticity fields, flow fields, and
pressure distribution on the wing surface
in the fling phase (t/T  0.55–0.65). (a)
Isovorticity surface plots of the wing; vor-
ticity is non-dimensionalized by U/c, and
the magnitude of the non-dimensional vor-
ticity is 2. (b) Non-dimensional flow veloc-
ity vector plots at section 0.7R (R is the
wing length) at the corresponding times.
(c) Pressure distributions on the upper
surface of the wing (first row) and the
lower surface of the wing (second row) at
the corresponding times.
33, 021905-8
 Physics of Fluids
FIG. 13. Vorticity fields and pressure distribution on the wing surface in the fling phase (t/T  0.55–0.65) and the subsequent downward/forward-sweeping phase (t/T
 0.65–1). (a) Non-dimensional isovorticity surface plots (top view) at t/T ¼ 0.65, 0.70, 0.75, and 0.8; vorticity is non-dimensionalized by U/cm, where cm is the mean chord
length of the wing. (b) Non-dimensional spanwise vorticity contours at the corresponding times. (c) Non-dimensional pressure distributions on the upper surface of the wing
(first row) and the lower surface of the wing (second row) at the corresponding times.
flapping cycle) can also be calculated. Dividing the mean power by
the mean vertical force gives the specific power, i.e., power used for
per Newton weight lifted, denoted by P*. For the tiny wasps in for-
ward flight, P* ranges from 2.18 W/N to 3.05 W/N; the average P*
for the five wasps is 2.56 W/N. For comparison, we also computed
the values of P* for the wasps in hover flight, studied by the authors
previously.26 For the eight wasps in hover flight,26 P* ranges from
2.08 W/N to 2.52 W/N; the average P* for the eight wasps in hover
flight is 2.36 W/N. We see that at medium speed forward flight, the
power requirement is slightly larger than that in hovering flight,

9.5% larger.
In order to achieve the large aerodynamic forces, as aforemen-
tioned, the tiny wasps adopt four different phases within one flapping
cycle (impulsive rowing, closing/upward-moving, fling, and down-
ward/forward sweeping). It is of great interest to know whether these
four stages are unique to Encarsia formosa in forward flight or they are
a strategy that is widely used by other miniature insects in a relatively
low-Re regime in forward flight. This will be investigated in our future
experiments.
IV. CONCLUDING REMARKS
The wings of miniature insects operate at very low Reynolds
number (Re in the order of 10), and the viscous effect is large. If their
wings flap like those of larger insects (Re in the order of 100–1000),
aerodynamic forces required for weight supporting and propulsion
cannot be produced. A new flapping mode must be used to overcome
the large viscous effect. In the present paper, we studied the wing
Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911
Published under license by AIP Publishing
ARTICLE scitation.org/journal/phf
motion pattern and aerodynamic force generation of a very small
wasp, Encarsia formosa, in forward flight. Our findings are the
following.
The flapping cycle consists of four phases: impulsive rowing (the
wings fast accelerate backward at very large angle of attack); clos-
ing/upward-moving (two wings come together at the back of the
insect and move upward); fling (the wings fling apart like opening a
book); and downward/forward sweeping. The first and second
phases are in the upstroke, and the third and fourth phases are in
the downstroke. In the impulsive rowing phase (the first half of the
upstroke), the wing smashes in the air and generates strong drag
that points forward and slightly upward, providing a large thrust
peak and a small vertical force peak. In the closing/upward-moving
phase (the second half of the upstroke), the wing slices through the
air and generates much smaller drag, giving a small negative verti-
cal force and negative thrust. In the fling phase (the start of the
downstroke), large drag is also produced by fast pushing the air,
providing some vertical forces and thrust; at the same time, a strong
leading edge vortex (LEV) is created on each wing. In the down-
ward/forward sweeping phase (the second, longer part of the down-
stroke), the wing carries the LEV on its upper surface and produces
large drag that points upward and backward, giving a large vertical
force and negative thrust.
The vertical force produced in the impulsive rowing, the fling,
and the downward/forward sweeping provides the weight supporting
force; the large thrust produced by the impulsive rowing and the fling
overcomes the body drag and the negative thrust produced by the
wings in the rest of the flapping cycle.
33, 021905-9
 Physics of Fluids ARTICLE scitation.org/journal/phf

The tiny insects overcome the strong viscous effect by fast
smashing the wings on the air in the first half of the upstroke, by
fast flinging the wings at the beginning of the downstroke, and by
carrying the LEV created at the fling to move in the rest of the
downstroke.
ACKNOWLEDGMENTS
This research was supported by grants from the National
Natural Science Foundation of China (Grant Nos. 11832004 and
11721202).

APPENDIX: DATA FOR OTHER FOUR WASPS

FIG. 14. Measured wing kinematics for

W2 (a), W3 (b), W4 (c), and W5 (d)
(mean 6 s.d.). For more details, see the
legend of Fig. 5.

Phys. Fluids 33, 021905 (2021); doi: 10.1063/5.0039911 33, 021905-10

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 Physics of Fluids ARTICLE scitation.org/journal/phf

DATA AVAILABILITY
The data that support the findings of this study are available
from the corresponding author upon request.

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