Stable hooks: biomechanics of heteromorph ammonoids with U-shaped body chambers

Abstract

ABSTRACT The biomechanics of uncoiled heteromorph ammonoids with body chambers that terminate in U-shaped hooks (ancylocones) were investigated with virtual and physical models of Audouliceras renauxianum. Virtual models were used to compute the hydrostatic properties of this morphotype. Audouliceras has the capacity for neutral buoyancy and this suggests that other taxa with similar proportions had this ability as well. Hydrostatic stability gradually increases during ontogeny, coincident with the larger degree of uncoiling. The juvenile planispiral stage has a similar stability and apertural orientation to the extant Nautilus. The adult stage, however, undergoes an increase in stability by a factor of over 3, while assuming an upward-facing posture. Counterintuitively, the stage during the formation of the shaft (before the growth of the U-shaped hook) is oriented horizontally. This intermediate stage would have had poor horizontal mobility due to the positioning of the hyponome below the centre of mass. The juvenile planispiral stage and mature stage, however, would have been well suited to horizontal backward movement with minimal rocking. Ancylocones are generally thought of as quasiplanktic vertical migrants. Thus, their relative horizontal swimming ability has been largely disregarded. Experiments on 3D printed, neutrally buoyant physical models reveal that hydrodynamic drag is indeed larger compared to Nautilus. However, Audouliceras could reach similar maximum horizontal velocities depending on the available thrust. Sepia-like thrusts yield velocities similar to equivalently sized Nautilus (c. 15 cm/s), while Nautilus-like thrusts yield velocities not much lower (c. 11 cm/s). Due to the hydrostatic properties of the ancylocone, the adult model undergoes less rocking (±4.5°) during movement than Nautilus (±10°). The minimal hydrodynamic consequences for ancylocones suggest that stability, orientation and directional efficiency are key selective pressures for some heteromorph shells, which may have primarily served as hydrostatic devices.