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< Leonid I. Frantsevich >

Leonid Frantsevich


Arthropod Structure & Development 2004, 33: 221-235.

Abstract. Twenty modes of stereotyped righting motions were observed in 116 representative species of Coleoptera. Methods included cine and sterocine recording with further frame by frame analysis, stereogrammetry, inverse kinematic reconstruction of joint angles, stroboscopic photography, recording of electromyograms, 3D measurements of the articulations etc. The basic mode consists of a search phase, ending up with grasping the substrate. and a righting, overturning phase. Leg coordination within the search cycle differs from the walking cycle with respect to phasing of certain muscle groups. Search movements of all legs appear chaotic, but the tendency to move in antiphase is still present in adjacent ipsilateral and contralateral leg pairs. The system of leg coordination might be split: legs of one side might search, while contralateral legs walk, or fore and middle legs walk while hind legs search. Elaborated types of righting include somersaults with the aid of contralateral or diagonal legs, pitch on elytra, jumps with previous energy storage with the aid of unbending between thoracic segments (well-known for Elateridae) or quick folding of elytra (originally described in Histeridae). Righting in beetles is compared with righting modes known in locusts and cockroaches. Search in a righting beetle is directed dorsad, while a walking insect search for the ground downwards. Main righting modes were schematized for possible application to robotics.

Unrestrained insects were filmed with a 32-mm camera, KONVAS, equipped with a stereoheadpiece. Each frame is viewed as a stereopair: a walking Lethrus apterus with an accessory body-fixed coordinate cross on its back. 3D coordinates of the cross and leg ends are reconstructed using stereogrammetric methods. After filming, the cross together with the landmarks of the body axes and positions of the subcoxal joints were measured in a 3D measurement microscope.

Knowing kinematic characteristics of a leg chain (orientation of joint axes, length of podomers), one can set arbitrary values of joint angles and compute positions of leg ends. It is the problem of direct kinematics. The problem of inverse kinematics is as follows: 3D positions of leg ends given, one must reconstruct values of joint angles in all six legs, including joints which are not seen in a stereoframe [reference 1].


Cyclic change of joint angles in the hind leg of Lethrus apterus during the walking (W) and the righting search (R).
Abbreviations of joints: FT – femoro-tibial, CF – coxo-trochanteral (=femoral), TC – thoracico-coxal joint. Joint angles in °.

Diagonal somersault in Blaps halophila. Selected stereoframes [reference 2].

We used photography with high-rate stroboscopic illumination for
recording of swift events instead of filming.

Jump of the click beetle, Lacon murinus, at the height almost 20 cm.
Collage of several stroboscopic recordings, flashing rate 100 Hz.
The beetle had white spots painted on the elytra and abdomen.

Jumps of the histerid beetle, Atholus duodecimstriatus.
Stroboscopic recordings, flashing rate 30 Hz [reference 3].

Somersault in a rose-chafer, Cetonia aurata on the glass.

Another metod of 3D filming is filming with a mirror, inclined by 45°. A videocamera is tilted by 90°, thus the real position of the insect and its mirror image is in a Portrait format.


(All videos are in AVI format. Left-click on an image to view in a separate window
or right-click and choose "Save File" to download)


Modes of righting in beetles: A - Dorcadion fulvum, B - Cassida nebulosa, C - Emus hirtus, D-F - Epicometis hirta, G,H - Byctiscus betulae.
Points of grasp or rest are indicated by circles.

Symmetrical somersault in a bug.

Bugs often behave like beetles when righting.

Symmetrical somersault in a pentatomid bug, Erthesina fullo.

Diagonal somersault in Erthesina fullo (video)

Diagrams of righting modes

A, B - righting by symmetrical grasp, C - righting by symmetrical somersault,
D - righting using turn of the prothorax.
Center of mass is marked by an asterisk, fixed grasping points - by the filled circles, slipping ground points - by blank circles.


[1] Frantsevich L.I., Shumakova I.D., Mokrushov P.A. 1984. Stereocinegrammetric analysis of insect movements (in Russian). Zhurnal Obschey Biologii 45: 89-97.

[2] Frantsevich L.I., Shumakova I.D. 1983. Kinematics of righting in Coleoptera. 3. Kinematics of the diagonal somersault (in Russian). Vestnik Zoologii (Kiev) N 2: 8-12.

[3] Frantsevich L.I. 1981. The jump of the black beetle (Coleoptera, Histeridae). Zool. Jb. Anat. 106: 333-348.


Collection of films:

Courtship dances in a fly, Lispe spp.

Stick friction in a lantern fly, Lycorma delicatula

Arolium of a hornet, Vespa crabro

Indirect closing of elytra in a cockchafer, Melolontha

Righting kinematics in beetles (Insecta: Coleoptera)

Leg coordination during turning on an extremely narrow substrate in a bug, Mesocerus marginatus (Heteroptera, Coreidae)

Swimming in the Diving Wasp Prestwichia aquatica (Hymenoptera: Trichogrammatidae)

Kinematics of elytra in beetles

Indirect closing of elytra in various beetles

Double rotation of the opening (closing) elytra in beetles (Coleoptera)

Actuation and performance of the elytron-to-body articulation in a diving beetle


I. I. Schmalhausen Institute of Zoology, 2004-2009