Anatomy and ontogenetic changes of the facial and gular musculature of the tetra Astyanax brucutu: A remarkable case of adaptation to durophagy.

Astyanax brucutu is a peculiar species of Neotropical tetra endemic from the Chapada Diamantina, a large plateau in northeastern Brazil. Individuals of this species undergo a dramatic ontogenetic shift in their diet that is accompanied by equally remarkable changes in their feeding apparatus. Whereas juveniles of A. brucutu feed mostly on algae, adults feed almost exclusively on an endemic species of hydrobiid snail and other associated living organisms that inhabit their dead shells (including infaunal invertebrates and algae). Skeletal adaptations associated with this change in diet were previously reported, but until now, the changes in the musculature remained mostly unknown. The present paper describes the facial and gular muscles, as well as the buccal ligaments of A. brucutu in different life stages, and identifies the major ontogenetic changes in these systems associated with the diet shift in the species. Such changes primarily involve expansions of specific portions of the adductor mandibulae and associated tendons and ligaments that likely represent adaptations to increase the biting power necessary to crush copious amounts of shells ingested by larger individuals of A. brucutu. Those adaptations are absent in specimens of any size of Astyanax cf. fasciatus, a sympatric congener lacking durophagous feeding habits. Anatomical comparisons and landmark-based principal components analysis (PCA) suggest that most specializations to durophagy in A. brucutu arose by peramorphosis. We also found that several of the muscular specializations of adults of A. brucutu are paralleled in species of Creagrutus and Piabina, two other characid genera distantly related to Astyanax, but that also feed on hard food items.