Mechanical and elemental characterization of ant mandibles: consequences for bite mechanics.

Mandible morphology has an essential role in biting performance, but the mandible cuticle can have regional differences in its mechanical properties. The effects of such a heterogeneous distribution of cuticle material properties in the mandible responses to biting loading are still poorly explored in chewing insects. Here, we tested the mechanical properties of mandibles of the ant species Formica cunicularia by nanoindentation and investigated the effects of the cuticular variation in Young's modulus (E) under bite loading with finite-element analysis (FEA). The masticatory margin of the mandible, which interacts with the food, was the hardest and stiffest region. To unravel the origins of the mechanical property gradients, we characterized the elemental composition by energy-dispersive X-ray spectroscopy. The masticatory margin possessed high proportions of Cu and Zn. When incorporated into the FEA, variation in E effectively changed mandible stress patterns, leading to a relatively higher concentration of stresses in the stiffer mandibular regions and leaving the softer mandible blade with relatively lower stress. Our results demonstrated the relevance of cuticle E heterogeneity in mandibles under bite loading, suggesting that the accumulation of transition metals such as Cu and Zn has a relevant correlation with the mechanical characteristics in F. cunicularia mandibles.

Mechanical demands of bite in plane head shapes of ant (Hymenoptera: Formicidae) workers.

Food processing can exert significant evolutionary pressures on the morphological evolution of animal appendages. The ant genus Pheidole displays a remarkable degree of morphological differentiation and task specialization among its workers. Notably, there is considerable variation in head shape within worker subcastes of Pheidole, which could affect the stress patterns generated by bite-related muscle contraction. In this study, we use finite element analysis (FEA) to investigate the effect of the variation in head plane shape in stress patterns, while exploring the morphospace of Pheidole worker head shapes. We hypothesize that the plane head shapes of majors are optimized for dealing with stronger bites. Furthermore, we expect that plane head shapes at the edges of each morphospace would exhibit mechanical limitations that prevent further expansion of the occupied morphospace. We vectorized five head shapes for each Pheidole worker type located at the center and edges of the corresponding morphospaces. We conducted linear static FEA to analyze the stresses generated by mandibular closing muscle contraction. Our findings indicate that plane head shapes of majors exhibit signs of optimization to deal with stronger bites. Stresses are distinctly directed along the lateral margins of the head, following the direction of muscle contraction, whereas the stresses on the plane head shapes of minors tend to concentrate around the mandibular articulations. However, the comparatively higher stress levels observed on majors' plane head shapes suggest a demand for cuticular reinforcement, like increased cuticle thickness or sculpturing pattern. Our results align with the expectations regarding the main colony tasks performed by each worker subcaste, and we find evidence of biomechanical limitations on extreme plane head shapes for majors and minors.

Climate and body size have differential roles on melanism evolution across workers in a worldwide ant genus.

One of the main aspects associated with the diversity in animal colour is the variation in melanization levels. In ectotherms, melanism can be advantageous in aiding thermoregulation through heat absorption. Darker bodies may also serve as a shield from harmful UV-B radiation. Melanism may also confer protection against parasites and predators through improving immunity responses and camouflage in regions with high precipitation, with complex and shaded vegetations and greater diversity of pathogens and parasites. We studied melanism evolution in the globally distributed ant genus Pheidole under the pressures of temperature, UV-B radiation and precipitation, while considering the effects of body size and nest habit, traits that are commonly overlooked. More importantly, we account for worker caste polymorphism, which is marked by distinct roles and behaviours. We revealed for the first time distinct evolutionary trajectories for each worker subcaste. As expected, major workers from species inhabiting locations with lower temperatures and higher precipitation tend to be more melanised. Curiously, we show a slight trend where minor workers of larger species also tend to have darker bodies when inhabiting regions with higher precipitation. Lastly, we did not find evidence for the effects of UV-B radiation and nest habit in the lightness variation of workers. Our paper explores the evolution of ant melanization considering a marked ant worker polymorphism and a wide range of ecological factors. We discuss our findings under the light of the Thermal Melanism Hypothesis, the Photoprotection Hypothesis and the Gloger's Rule.

Imperfect synchrony in animal displays: why does it occur and what is the true role of leadership?

Synchrony can be defined as the precise coordination between independent individuals, and this behaviour is more enigmatic when it is imperfect. The traditional theoretical explanation for imperfect synchronous courtship is that it arises as a by-product of the competition between males to broadcast leading signals to attract female attention. This competition is considered an evolutionary stable strategy maintained through sexual selection. However, previous studies have revealed that leading signals are not honest indicators of male quality. We studied imperfect courtship synchrony in fiddler crabs to mainly test whether (i) signal leadership and rate are defined by male quality and (ii) signal leadership generates synchrony. Fiddler crab males wave their enlarged claws during courtship, and females prefer leading males-displaying ahead of their neighbour(s). We filmed groups of waving males in the field to detect how often individuals were leaders and if they engaged in synchrony. Overall, we found that courtship effort is not directly related to male size, a general proxy for quality. Contrary to the long-standing assumption, we also revealed that leadership is not directly related to group synchrony, but faster wave rate correlates with both leadership and synchrony. This article is part of the theme issue 'Synchrony and rhythm interaction: from the brain to behavioural ecology'.

Mandibular morphology, task specialization and bite mechanics in Pheidole ants (Hymenoptera: Formicidae).

Ants show remarkable ecological and evolutionary success due to their social life history and division of labour among colony members. In some lineages, the worker force became subdivided into morphologically distinct individuals (i.e. minor versus major workers), allowing for the differential performance of particular roles in the colony. However, the functional and ecological significance of these morphological differences are not well understood. Here, we applied finite element analysis (FEA) to explore the biomechanical differences between major and minor ant worker mandibles. Analyses were carried out on mandibles of two Pheidole species, a dimorphic ant genus. We tested whether major mandibles evolved to minimize stress when compared to minors using combinations of the apical tooth and masticatory margin bites under strike and pressure conditions. Majors performed better in pressure conditions yet, contrary to our expectations, minors performed better in strike bite scenarios. Moreover, we demonstrated that even small morphological differences in ant mandibles might lead to substantial differences in biomechanical responses to bite loading. These results also underscore the potential of FEA to uncover biomechanical consequences of morphological differences within and between ant workers.