The ptychoid defensive mechanism in Euphthiracaroidea (Acari: Oribatida): a comparison of muscular elements with functional considerations.

Ptychoidy is a mechanical predator defence in some groups of Oribatida (Acari), where the animals can retract their legs into the idiosoma and encapsulate. This mechanism is enabled by a number of morphological adaptations. We used the non-invasive technique of synchrotron X-ray microtomography to compare muscular elements involved in ptychoidy of two species from the Euphthiracaroidea (Oribotritia banksi and Rhysotritia ardua) which differ in a number of cuticular elements involved in ptychoidy. We hypothesize that a strong functional correlation exists between these cuticular structures and their corresponding musculature. We found a number of distinct differences concerning quantitative and qualitative muscle morphology. For testing the functional impact of different muscle configurations we simulated two conditions in silico (encapsulated / opened) and analysed the spatial relative force vectors of the prodorsum lateral adjustor muscles (pla) which are responsible for retraction and adjustment of the prodorsum during encapsulation. We show that the functional morphology of these muscles strongly differs between the two species and that this can be explained by the structure of corresponding cuticular elements. Furthermore, the dynamics of pla, as measured by the extent of contraction during encapsulation, is more than two times higher in R. ardua than in O. banksi.