Sniffing out Stingray Noses: The Functional Morphology of Batoid Olfaction.

Batoid fishes (rays, skates, sawfishes, and guitarfishes) are macrosmatic, meaning they rely on their sense of smell as one of the primary senses for survival and reproduction. Olfaction is important for long-distance tracking and navigation, predator and prey recognition, and conspecific signaling. However, the mechanisms by which batoids harness odorants is unknown. Without a direct pump-like system, it is hypothesized that batoids irrigate their nostrils via one or a combination of the following: the motion pump, buccopharyngeal pump, pressure (ex. pitot-like mechanism), or a shearing force (ex. viscous entrainment). These mechanisms rely on the size, shape, and position of the nostrils with respect to the head and to each other. Batoids are united as a group by their dorsoventrally compressed body plans, with nostrils on the ventral side of their body. This position presents several challenges for odor capture and likely limits the effectivity of the motion pump. Batoid fishes display an expansive nasal morphology, with inlet nostrils ranging from thin, vertical slits to wide, horizontal ovals to protruding, tube-like funnels, and more. In this paper, a morphometric model is developed to quantify the vast diversity in batoid nose shapes, sizes, and positions on the head in an ecological and functional framework. Specifically, swimming mode, lifestyle, habitat, and diet are examined for correlations with observed nasal morphotypes. Morphometric measurements were taken on all 4 orders present in Batoidea to broadly encompass batoid nasal diversity (Rhinopristiformes 4/5 families; Rajiformes 2/4 families; Torpediniformes 4/4 families; Myliobatiformes 8/11 families). All batoid external nasal diversity was found to be categorized into 5 major morphological groups and were termed: flush nare [circle, comma, intermediate], open nare, and protruding nare. Several morphometric traits remained significant when accounting for shared ancestry, including the position and angle of the nostril on the head, the width of the inlet hole, and the spacing of the nostrils from each other. These measurements were found to be closely correlated and statistically significant with the swimming mode of the animal. This study provides the first crucial step in understanding batoid olfaction, by understanding the diversity of the morphology of the system. Because odor capture is a strictly hydrodynamic process, it may be that factors relating more directly to the fluid dynamics (i.e., swimming mode, velocity, Reynolds number) may be more important in shaping the evolution of the diversity of batoid noses than other ecological factors like habitat and diet.