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The centroid size of each wing was calculated by determining the relative position (landmark) of 12 vein intersections [after 53] and then calculating the square root of the sum of squared distances between each landmark and the centroid of each forewing [54].
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Torsional stiffness of the wing and the distribution of mass along the wing are calculated.
The camber of the mid wing is calculated as the perpendicular distance from the chord (wrist P V t [pt4 13] to the point P V i [pt 15] divided by the length of the chord.
PHA (Sigma L-8754) in phosphate buffered saline (PBS; 0.45 µg in 50 µl [30], [31] was injected into both wings webs of each bird. The mean response of both wings was calculated and used in all analyses.
This parameter can be defined as a quantitative measure of wing area distribution along the wing axis [ 10] and for a pair of wings is calculated as follows: r ^ 2 S = ∫ 0 1 c ^ r ^ d r ^, where ĉ is the normalized wing chord and r ^ is the non-dimensional radius equal to r/ R, where r is the distance to the wing base on the chord c and R is wing length.
Wing loading was calculated as total body weight / total wing area (cf. Betts and Wootton 1988; Breuker et al. 2010).
The abnormal-wing-posture penetrance was calculated as the percentage of flies with either held-up or drooped wing posture.
A template consisting of the wing outline and veins L2 to L5 is fitted to the wings, and then wing length, wing width, and wing area are calculated.
Aspect ratio, which is usually used to model the overall shape of the wing (e.g. Rayner 1988), was calculated as (wingspan)/wing area.
Using these parameters the mean chord c = S/2 b, the non-dimensional aspect ratio AR = 4 b/ S and the wing loading Q = mg/ S was calculated, where m is the body mass and g the gravitational acceleration (Table 1).
During flapping we could not distinguish heart beats from wing beats, and therefore fH during flapping flight was calculated from the first spikes that followed a series of wing beats during glides.
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