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A Biological Screw in a Beetles Leg One possible disadvantage, the danger of over-
winding the thread, then is negligible, because the
depressor muscles are already fully contracted at
Thomas van de Kamp,1,2,3 Patrik Vagovi,1 Tilo Baumbach,1 Alexander Riedel2* that point. We suggest that an advantage of this
construction is that the leg comes to a stable resting
n many animals, movement relies on joints an apophysis differing between the fore, mid, and position, preventing passive straining of leg muscles,

I between skeletal features, analogous to joints hind legs. The maximum rotation achieved by the
between moving parts of mechanical machines. muscles is 90 (protrochanter) and 130 (meso-
Some, such as ball-and-socket joints, are readily and metatrochanter), respectively. The rotational
which would not be accomplished by an axle con-
struction. If the weevil's leg is pulled in the op-
posite direction over its back, the muscles are torn
found in both living organism and mechanical axes of the trochanters are positioned at approx- off, and the trochanter can be neatly dislodged from
engineering (1), whereas others are restricted to one imate right angles to the femoral axes. The overall the coxal nut. Such dislocation can be produced
of these realms. We report the finding of a func- translation along the axes of rotation ranges from with the help of delicate forceps. Under natural cir-
tional screw-and-nut system in the coxa-trochanteral 31.5 m (0.35 m/) to 65.0 m (0.50 m/). The cumstances, this is a highly unlikely event, because
joints of the legs of the Papuan weevil Trigonopter- diameter of the threaded part of the trochanter the beetles natural defensive reflex is to bring the
us oblongus (Pascoe) (2). We reveal the systems decreases toward its proximal end. legs into a ventral position if attacked by a predator.
structure and function through interactive three- Our preliminary microscopic examination of The presumably primitive coxa-trochanteral
dimensional (3D) reconstructions created from other weevil genera [listed in (6)] showed sim- joint in ground beetles (Carabidae) possesses a sym-
synchrotron-based x-ray microtomography (CT) ilarities with the coxa-trochanteral joint as revealed metrical pair of condyles and works as a hinge. In
(3) of museum specimens. These reconstructions here for Trigonopterus. The screw-and-nut system Chrysomeloidea and in Curculionoidea, one con-

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allow the simulation of movements in this artic- appears to be widespread among weevils and may dyle is reduced (in the foreleg, the posterior condyle;
ulation (Fig. 1 and figs. S3 to S5). indeed represent a basic character of the family. In in the mid and hind legs, the anterior condyle) and
The three major joints of an insect leg, the coxa- other families of beetles, coxa-trochanteral joints the other is produced into a thorn. These joints pos-
trochanteral, trochantero-femoral and femoro-tibial that perform a rotation accompanied by a transla- sess a greater dorsoventral mobility, which is better
articulations, are usually considered as hinges (4). tory movement along the axis have been observed suited for a life on twigs and foliage. Many weevil
However, in the species of hyperdiverse Trigonop- in Scarabaeoidea (7) and Chrysomelidae (3, 8). species depend on tibial spines to provide strong
terus weevils studied here, the coxa-trochanteral However, their constructions [supporting online foothold while the rostrum is pushed into the sub-
joint is highly modified and allows rotational move- material (SOM) text] are markedly different. A strate. Possibly, the forces acting on the leg joints
ment combined with a single-axis translation. continuous screw thread is either absent (in Scara- during this feeding process unique to weevils have
The pro- and mesocoxae of T. oblongus are baeoidea) or incomplete (a spiral ridge of trochan- driven the transformation from the chrysomeloid
subspherical in shape, the metacoxae bulging ter covers 210 and the inner thread of the coxa joint type to the more robust one in weevils.
somewhat forward. Their apical opening (fig. S1, 180 in the chrysomelid genus Cryptocephalus); in-
References and Notes
A and B) is wide and circular and mesially shows stead, a process of the coxa is interlocked with a 1.G. Bgelsack et al., Theory Biosci. 119, 104 (2000).
a notch marking the start of a well-defined inner pit in the trochanter. This mechanism, securing the 2.A. Riedel, D. Daawia, M. Balke, Zool. Scr. 39, 63 (2010).
thread that continues internally for 345. Thus, joint, has disappeared in the weevils examined. 3.O. Betz et al., J. Microsc. 227, 51 (2007).
the apical portions of the coxae closely resemble It is remarkable that in the case of the weevil 4.L. Frantsevich, W. Wang, Arthropod Struct. Dev. 38, 16
engineered screw nuts. The trochanters (fig. S1, leg a rotary movement is accomplished by a screw-
5. O. Larsn, Opusc. Entomol. 30 (suppl.), 1 (1966).
C and D) of all of the weevils legs are similar, and-nut system. In engineering, such systems are 6. Materials and methods are available as supporting
carry external spiral threads that cover 410, and mainly used for fixing connections, whereas an material on Science Online.
are perfectly compatible with the threads in the axle would be used for a simple rotation. The pos- 7. F. Reuleaux, in Lehrbuch der Kinematik, F. Reuleaux, Ed.
respective coxal openings. Proximally, each tro- sible advantages and disadvantages of a screw- (Friedrich Vieweg und Sohn, Braunschweig, Germany, 1900),
vol. 2, pp. 723777.
chanter is produced into a thorn that spears the and-nut system over an axle in a leg joint may be 8. M. L. Chamorro-Lacayo, A. S. Konstantinov, Zootaxa 676,
entire coxa; when the hind leg is depressed, the most apparent at its end points. In ventral position, 1 (2004).
thorns tip penetrates the opposite coxal wall when fully tightened, the joint comes to a dead stop. Acknowledgments: We are grateful to F. Glaw encouraging
through a small opening, thus us to prepare the initial manuscript, to S. Scharf for
performing the segmentation of CT scans, and to
stabilizing the joint along D. Pelliccia for his assistance during the scans. M. Balke,
the rotational axis and se- O. Betz, and M. Spies kindly revised earlier drafts. The
curing it against jamming. specimens examined are deposited at SMNK. The image data
Three muscles attach to are stored at www.morphdbase.de?A_Riedel_20110523-M-4.1.
This work was supported by grant RI 1817/3-1 from the
the trochanter between the German Research Foundation.
screw thread and the prox-
imal thorn. The composition Supporting Online Material
of muscles (fig. S2) con- www.sciencemag.org/cgi/content/full/333/6038/52/DC1
Materials and Methods
forms with that previously SOM Text
described for weevils and Figs. S1 to S5
most other polyphagan bee- References
tles (5). Outward rotation is 14 February 2011; accepted 13 May 2011
accomplished by a muscle 10.1126/science.1204245
that originates at the coxal Fig. 1. The 3D reconstruction [(A) to (E)] of coxa (green) and trochanter
wall and attaches directly to (yellow) of left hind leg of T. oblongus. (A) Depressed position. (B) Elevated ANKA - Institute for Synchrotron Radiation, Karlsruhe Institute
the trochanter; inward rota- position. (C) Coxa cut horizontally along rotation axis; dorsal aspect of tro- of Technology, D-76344 Eggenstein-Leopoldshafen, Germany.
State Museum of Natural History (SMNK), D-76133 Karlsruhe,
tion is by two muscles attach- chanter while leg depressed. (D). Isolated trochanter showing external spiral Germany. 3Institute for Zoology II, Heinrich Heine University,
ing to the trochanter via a thread and tendon (t). (E) Dorsal portion of coxa corresponding to ventral D-40225 Dsseldorf, Germany.
tendon, one originating from portion of (C). (F) Scanning electron microscope photograph of the right *To whom correspondence should be addressed. E-mail:
the coxal wall, the other from metatrochanter showing posterior condyle (c) and external spiral thread (e). riedel@smnk.de

52 1 JULY 2011 VOL 333 SCIENCE www.sciencemag.org

A Biological Screw in a Beetle's Leg
Thomas van de Kamp, Patrik Vagovic, Tilo Baumbach and
Alexander Riedel (June 30, 2011)
Science 333 (6038), 52. [doi: 10.1126/science.1204245]

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