Ammonite aptychi: Functions and role in propulsion

Seven previous proposals of aptychus (sensu stricto) function are reviewed: lower mandible, protection of gonads of females, protective operculum, ballasting, flushing benthic prey, filtering microfauna and pump for jet propulsion. An eighth is introduced: aptychi functioned to actively stabilize the rocking produced by the pulsating jet during forward foraging and backward swimming. Experiments with in-air models suggest that planispiral ammonites could lower their aperture by the forward shift of a mobile cephalic complex. In the experiments, the ventral part of the peristome is lowered from the lateral resting (neutral) position by the added “ballast” of a relatively thin Laevaptychus to an angle < 25° from horizontal with adequate stability to withstand the counter-force produced by the jet of the recurved hyponome. However, of the shell forms tested, only brevidomes with thick aptychi, e.g., the Upper Jurassic Aspidoceratidae with Laevaptychus and average whorl expansion rates, were stable enough to swim forward by jet propulsion at about Nautilus speed (∼ 25 cm/s). We propose that aptychus function most commonly combined feeding (jaw, flushing, filtering) with protection (operculum), and, more rarely, with locomotion (ballast, pump, diving and stabilizing plane). Aptychi may thus have been multi-functional.