Finite Element Models based on MicroCT data show

which stag beetle species are most vulnerable to

break their jaws in battles

J. Goyens1,2, J. Dirckx2, P. Aerts1,3

1
University of Antwerp, Laboratory of Functional Morphology, Universiteitsplein 1, B-2610
Antwerp, Belgium,
2
University of Antwerp, Laboratory of BioMedical Physics, Groenenborgerlaan 171, B-2020
Antwerp, Belgium
3
University of Ghent, Department of Movement and Sport Sciences, Watersportlaan 2, 9000
Ghent, Belgium

Aims
Through sexual selection, male stag beetles have evolved
extremely long and ornamented jaws. In spectacular battles,
they use these jaws to bite opponents forcefully, in order to
gain mating rights (see Fig. 1)1. Therefore, stag beetle jaws
must be constructed robustly to prevent breaking2. However,
at the same time, they must be as lightweight as possible to
minimize the costs of walking and flying3,4.
Species in the stag beetle family are known for the large
variation in sizes and shapes of their jaws. They also utilize
their jaws is diverse ways, and their bite force differs as well.
Therefore, different species may use different strategies to
prevent breaking their jaws. We investigated the robustness
of the jaws of 16 stag beetle species with Finite Element
Models (FE-models) that are based on MicroCT scans.

Figure 1: Fighting male Cyclommatus metallifer stag beetles

Method
We made microCT scans of the heads of 16 different species with a Skyscan 1172 high
resolution micro CT scanner (Bruker microCT, Kontich, Belgium; see Table 1). We used these
microCT scans for 2 goals: to determine the muscle force of the bite muscles and to create a
3D model of the jaws.

Bite muscle force The force of a stag beetle muscle is determined by the area of its
attachment on the head capsule. However, this was often not visible anymore on the microCT
scans of our dried samples, and we therefore used the complete area of the head surface as
a proxy. We calculated this area in the 3D image processing software Amira (Amira 5.4.4; 64-
bit version, FEI, Hillsboro, OR, USA).

3D jaw models In the same software package, we designated which voxels belong to the jaw
exoskeleton (cuticle) with a combination of grey-scale thresholding and manual corrections in
the three orthogonal views. Next, we created a volume mesh, consisting of tetrahedra with
TetGen software5.

FE models Three things are required to construct the FE models: a 3D model, material
properties and mechanical constraints. Further, we imported the 3D jaw models in FEBio
1.4.16. We applied material properties that are characteristic for stag beetle cuticle: a Youngs
modulus of 5.1 GPa and a Poisson ratio of 0.3. Finally, we added mechanical constraints to
the 3D jaw models (see Fig. 2): an force pulled at the muscle attachment location, the jaw
hinge was allowed to rotate and the jaw tip was prohibited from moving in x direction (which
mimics the body of the opponent that is being bitten).

Table 1: Micro CT scan parameters.

Voxel size Voltage Current Rotation Frame
Nr Species
(m) (kV) (A) angle averaging
1 P.giraffa 13.44 50 198 0.4 4
2 P. mohnikei 13.44 50 198 0.4 3
3 H. parryi 13.44 50 198 0.4 4
4 D. parryi 13.44 60 165 0.3 3
5 D. bucephalus 13.44 60 165 0.3 3
6 D. titanus 13.44 49 200 0.4 3
7 D. alcides 13.44 49 200 0.3 3
8 C. metallifer 8.2 70 141 0.2 5
9 L. cervus 13.44 56 177 0.4 3
10 L. adophinae 13.44 50 198 0.35 4
11 P. davidisi 13.44 49 200 0.25 5
12 N. obesus 10.75 49 200 0.35 4
13 C. lucifer 13.44 49 200 0.35 4
14 P. bison 13.44 49 200 0.35 4
15 P. oberthuri 13.44 49 200 0.25 5
16 P. senegalensis 13.44 50 198 0.3 5

Results and Discussion

Figure 2 shows the material stress in the jaws of the 16 stag beetle species when they bite
with their natural muscle force (as calculated from their head size). The species differ in the
location of the maximal material stress. Hence, they would break at a different location if they
would exceed their ultimate stress. In some species, only a small part of their jaw is prone to
the highest stress (e.g. species nr 11 and 12). In other species, the highest stress is not
concentrated in a spot (e.g. species nr 4), which means that their jaw is more equally robust
along its length. Further, also the magnitude of the highest material stress differs between
species (see Fig. 2 and 3A). Species nr 10 has a very low maximal material stress, while
species nr 3 and 8 have a very high maximal material stress, even though they have an
average muscle force. The other species have a remarkable similar maximal material stress,
even though they cover a large range of muscle forces.
Species that bite more forcefully, may also invest in more jaw material to make their jaws more
robust. To investigate this hypothesis, we scaled the input force in the FE models according to
the exoskeletal volume (see Fig. 3B). These models simulate the hypothetical situation where
all jaws would have an equal amount of material (same volume models). However, the
hypothesis is not completely confirmed by our results: the different species not have the same
maximal material stress after scaling for jaw volume. Rather, species with heavier jaws (more
jaw volume) show higher material stresses. Further investigations will show what causes this
effect. For example, species that invest in more jaw volume, may not only do so to increase
their jaw robustness, but they may also (partially) use this additional volume to increase their
jaw length. Indeed, behavioral investigations have shown that longer stag beetle jaws are
advantageous in battles because they enable a further reach towards opponents 7.
 Figure 2: Comparison of the material stress between stag beetle species. For visualization
purposes, the jaws are depicted with the same jaw length. On the model of the first species,
the mechanical constraints are shown in purple.
 Figure 3: (A) Relation between the bite muscle force of the species and the Von Mises stress
in FE models with the bite muscle force as input force. (B) Relation between the jaw volume of
the species and the Von Mises stress in same volume FE models.

Conclusion
Stag beetle species differ largely in the shape, length and robustness of their jaws. They also
exert a different muscle force on their jaws. Hence, they probably use their jaws in different
ways and for different purposes. Despite the differences in their morphology and behavioural
strategies, our FE analyses show that (except for a few exceptions) the species undergo
remarkable similar material stresses in their jaws. This probably implies that species that
perform more perilous activities with their jaws, have adapted their jaw robustness accordingly
to keep the risk of failure at the same level.

References:
1. Kawano, K., Sexual dimorphism and the making of oversized male characters in
beetles (Coleoptera) Annals of Biomedical Engineering 99, 327-341, 2006
2. Goyens, J., Soons, J., Aerts, P. and Dirckx, J., Finite Element modelling reveals
force modulation of jaw adductors in stag beetles Journal of the Royal Society
Interface 11, 20140908, 2014
3. Goyens, J., Dirckx J. and Aerts P., Costly sexual dimorphism in Cyclommatus
metallifer stag beetles Functional Ecology 29, 35-43, 2015
4. Goyens, J., Van Wassenbergh, S., Dirckx J. and Aerts P., Cost of Flight and the
Evolution of Stag Beetle Weaponry (In Revision) Journal of the Royal Society
Interface
5. Si, H., TetGen: a quality tetrahedral mesh generator and thre-dimensional
Delaunay triangulator, Users manual, v. 1.4 tetgen.org 2006
6. Maas, S., Ellis, B., Ateshian, G., Weiss, J., FEBio: Finite Elements for
Biomechanics Journal of Biomechanical Engineering 134, 011005, 2012
7. Goyens, J., Dirckx J. and Aerts P., Stag beetle battle behaviour and its associated
anatomical adaptations (In Revision) Journal of Insect Behavior