Comparative morphology of the rhinarium and upper lip in sigmodontine rodents: Refined nomenclature, intertribal variation in a phylogenetic framework, and functional inferences

Rodents have received substantial attention in the study of olfaction. However, the rhinarium, the naked part of the nose, which plays an important role in chemical, tactile, and thermal perception, has been relatively overlooked. This study presents a comprehensive analysis of the rhinarium morphology and spatially associated structures (i.e., upper lip, and philtrum) in sigmodontines, a diverse group within the Cricetidae rodents. The research covers 483 specimens representing 145 species, accounting for 74% of genera in the clade, including all 13 recognized tribes, three incertae sedis genera, and the murid representatives Mus musculus and Rattus norvegicus. The inconsistent use of terminology in describing rhinarium traits across the literature poses a challenge for comparative analyzes. To address this issue, a standardized terminology was proposed to characterize the rhinarium. A paired complex protuberance typically with epidermal ridges (i.e., rhinoglyphics), termed here the tubercle of Hill, was identified as a distinctive feature in muroid rhinaria. Comparative assessments among tribes revealed unique sets of features defining each major clade, encompassing variations in hairiness, dorsum nasi complexity, size and positioning of the tubercle of Hill, and other key attributes. Two primary rhinarium configurations were discerned: one shared by Oryzomyalia and Sigmodontini and another specific to Ichthyomyini. The former groups display a ventrally positioned rhinarium prominently featuring the tubercle of Hill and sculptured areola circularis. In contrast, Ichthyomyini exhibit a frontally directed rhinarium characterized by an enlarged dorsum nasi fused to the tubercle of Hill, resulting in a distinctive “cherry” appearance. Convergent rhinarium structures observed in fossorial species, characterized by well-developed plica alaris and hair fringes, are presumed to mitigate potential damage during digging. Conversely, semiaquatic carnivorous sigmodontines showcase an integrated apical structure in their rhinarium, facilitating enhanced somatosensory capabilities crucial for predation activities during diving expeditions.