Syn vivo hydrostatic and hydrodynamic properties of scaphitid ammonoids from the U.S. Western Interior

Abstract

Scaphitid ammonoids were ubiquitous and significant components of the Western Interior Seaway during the Late Cretaceous. This group is characterized by a recurved hook at maturity that deviates from the juvenile whorls. Such a modification seems counterproductive to active locomotion and to manage a biologically effective orientation that facilitates efficient feeding and swimming. Virtually reconstructed 3D hydrostatic models reveal that the examined mature scaphitids had the capacity for neutral buoyancy while assuming a stable, upward-facing orientation in the water column during life. Models of juvenile Hoploscaphites nicolletii suggest that scaphitid apertures were oriented only slightly more horizontal than adults. The hydrostatic influence of sexual dimorphism was explored with the species Hoploscaphitescrassus. The inflated macroconch has a lower stability and higher hydrodynamic drag compared to its microconch counterpart. The effect of shell compression was investigated by comparing H. crassus and the more compressed H. nicolletii. The latter species has a relatively high stability and much less hydrodynamic drag during movement. The mature U-shaped body chamber distributes organismal mass in a way that increases stability, and simultaneously orients the soft body so that propulsive energy is efficiently transmitted into horizontal backwards movement with minimal rocking. Swimming velocities computed from hydrodynamic drag experiments suggest that scaphitids were relatively slow swimmers with compressed forms attaining slightly higher velocities (when scaled by mass). Hydrodynamic lift was investigated with computational fluid dynamics simulations. These experiments revealed that the overall shape of the shell is responsible for significant lift in the upwards direction, which is not heavily influenced by ornamentation. This explains how a reduced soft body can overcome and manage a slightly negatively buoyant condition during life. Therefore, the seemingly cumbersome shape and orientation of the scaphitid morphotype may not have been a hindrance during locomotion.