| 1. | The movements of alae and elytra are determined. Both move with the same frequency and in the same phase, but the range of the elytral stroke never reaches below the frontal plane. In downstroke the elytra are at an angle positive to the airstream (horizontal flight). |
| 2. | After exact investigation of the construction of the elytra (depth, width, length, surface, torsion and velocity of each point of the elytra) the most effective point in the production of lift during the downstroke was determined. |
| 3. | After loss of one third of the surface of the elytra the beetles cannot fly horizontally. In tethered flight the wing stroke frequency of the alae does not increase when the elytra are shortened. Shortening of the elytra does not interfere with the resonance system of the pterothorax. |
| 4. | The lift of the elytra results from active and passive components (more than 17% of the body weight). In the airstream (front 2.00 m/s, average body weight 944 mp) the passive lift is 14.3% for elytra in the down position; 3.1% of the lift is caused by the active downstroke. |
| 5. | The elytra of these beetles are indispensible for horizontal flight. Primarily they produce lift passively with added help from the downstroke, and have no function for the drag. The effect of upstroke is discussed. |
