Wonderfully weird: the head anatomy of the armadillo ant, Tatuidris tatusia (Hymenoptera: Formicidae: Agroecomyrmecinae), with evolutionary implications

Tatuidris tatusia BROWN & KEMPF, 1968, the armadillo ant, is a morphologically unique species found in low to high elevation forests in regions of Central and South America. It is one of only two extant representatives of the subfamily Agroecomyrmecinae, and very little is known about the biology of these ants, which are almost exclusively collected from leaf litter and have rarely been seen alive. Here, we illuminate the functional morphology and evolution of this species via detailed anatomical documentation of their exceptionally modified head. We describe and illustrate the skeletomuscular system, digestive tract, and cephalic glands based on high-resolution micro-computed tomography scan data. We hypothesize that the modifications which produce the unusual "shield-like" head shape are the result of complex optimizations for mandibular power, physical protection, and balance. The most conspicuous cephalic features are the broadening of the frontal region and foreshortening of the postgenal region. The former characteristic is likely also associated with the lateral position of the antennal scrobe, the inverted antennal articulation, and the broad attachment surface for the mandibular adductor muscles. This head geometry also comes with a degree of internal restructuring of the tentorium and the antennal musculature, which have a unique configuration among ants studied so far. The mandibular blades, and their articulations and muscles, are highly distinctive compared with previously evaluated species. Using a 3D-printed model, we were able to hypothesize their entire range of motion as the mandibles fit tightly into the oral foramen. Finally, we compare T. tatusia across other related subfamilies and discuss the evolution of the Agroecomyrmecinae and other species-poor and phylogenetically isolated "relictual" lineages.