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Externally shelled cephalopods with coiled, planispiral conchs were ecologically successful for hundreds of millions of years. These animals displayed remarkable morphological disparity, reflecting comparable differences in physical properties that would have constrained their life habits and ecological roles. To investigate these constraints, self-propelling, neutrally buoyant, biomimetic robots were 3D-printed for four disparate morphologies. These robots were engineered to assume orientations computed from virtual hydrostatic simulations while producingNautilus-like thrusts. Compressed morphotypes had improved hydrodynamic stability (coasting efficiency) and experienced lower drag while jetting backwards. However, inflated morphotypes had improved maneuverability while rotating about the vertical axis. These differences highlight an inescapable physical tradeoff between hydrodynamic stability and yaw maneuverability, illuminating different functional advantages and life-habit constraints across the cephalopod morphospace. This tradeoff reveals there is no single optimum conch morphology, and elucidates the success and iterative evolution of disparate morphologies through deep time, including non-streamlined forms.

 

Abstract The internal architecture of chambered ammonoid conchs profoundly increased in complexity through geologic time, but the adaptive value of these structures is disputed. Specifically, these cephalopods developed fractal-like folds along the edges of their internal divider walls (septa). Traditionally, functional explanations for septal complexity have largely focused on biomechanical stress resistance. However, the impact of these structures on buoyancy manipulation deserves fresh scrutiny. We propose increased septal complexity conveyed comparable shifts in fluid retention capacity within each chamber. We test this interpretation by measuring the liquid retained by septa, and within entire chambers, in several 3D-printed cephalopod shell archetypes, treated with (and without) biomimetic hydrophilic coatings. Results show that surface tension regulates water retention capacity in the chambers, which positively scales with septal complexity and membrane capillarity, and negatively scales with size. A greater capacity for liquid retention in ammonoids may have improved buoyancy regulation, or compensated for mass changes during life. Increased liquid retention in our experiments demonstrate an increase in areas of greater surface tension potential, supporting improved chamber refilling. These findings support interpretations that ammonoids with complex sutures may have had more active buoyancy regulation compared to other groups of ectocochleate cephalopods. Overall, the relationship between septal complexity and liquid retention capacity through surface tension presents a robust yet simple functional explanation for the mechanisms driving this global biotic pattern. 

Measuring locomotion tactics available to ancient sea animals can link functional morphology with evolution and ecology over geologic timescales. Externally-shelled cephalopods are particularly important for their central roles in marine trophic exchanges, but most fossil taxa lack sufficient modern analogues for comparison. In particular, phylogenetically diverse cephalopods produced orthoconic conchs (straight shells) repeatedly through time. Persistent re-evolution of this morphotype suggests that it possesses adaptive value. Practical lateral propulsion is ruled out as an adaptive driver among orthoconic cephalopods due to the stable, vertical orientations of taxa lacking sufficient counterweights. However, this constraint grants the possibility of rapid (or at least efficient) vertical propulsion. We experiment with this form of movement using 3D-printed models of Baculites compressus, weighted to mimic hydrostatic properties inferred by virtual models. Furthermore, model buoyancy was manipulated to impart simulated thrust within four independent scenarios (Nautilus-like cruising thrust; a similar thrust scaled by the mantle cavity of Sepia; sustained peak Nautilus-like thrust; and passive, slightly negative buoyancy). Each model was monitored underwater with two submerged cameras as they rose/fell over ~2 m, and their kinematics were computed with 3D motion tracking. Our results demonstrate that orthocones require very low input thrust for high output in movement and velocity. With Nautilus-like peak thrust, the model reaches velocities of 1.2 m/s (2.1 body lengths per second) within one second starting from a static initial condition. While cephalopods with orthoconic conchs likely assumed a variety of life habits, these experiments illuminate some first-order constraints. Low hydrodynamic drag inferred by vertical displacement suggests that vertical migration would incur very low metabolic cost. While these cephalopods likely assumed low energy lifestyles day-to-day, they may have had a fighting chance to escape from larger, faster predators by performing quick, upward dodges. The current experiments suggest that orthocones sacrifice horizontal mobility and maneuverability in exchange for highly streamlined, vertically-stable, upwardly-motile conchs. 

Of the many shell morphologies produced by ammonoid cephalopods, the helical torticone shape appears poorly-suited to rapid locomotion. We investigate torticone hydrostatics and hydrodynamics through virtual modeling, computational fluid dynamics simulations, and water-chamber experiments, using the Cenomanian (Cretaceous) turrilitid Mariella brazoensis (Roemer, 1852) as a test case. Our hydrostatic model suggests that M. brazoensis, and other torticones, could attain neutral buoyancy. This morphotype is highly stable compared to planispiral cephalopods with a slightly tilted, apex-upward orientation. The corresponding mass distribution, relative to the source of jet propulsion at the hyponome, suggests that jet thrust would be more efficiently transmitted into upwards movement than horizontal movement. Most directions of thrust would send the shell spinning about its vertical axis. We 3D printed shell models to have either surpluses or deficiencies in buoyancy that imparted estimated thrusts of extant cephalopod analogues in the vertical directions within a water chamber. The models consistently rotate aperture-backwards during upward movement, and aperture-forwards during downward movement. A neutrally buoyant model was used to assess rotational aptitude during active locomotion. The model required low torques to sustain rotation. Simulations of water flow around the shell support the movement directions observed in the physical experiments and demonstrate that hydrodynamic drag is lower in the vertical directions than the horizontal directions. These results show that the animal within a torticone shell could spin about its vertical axis easily; perhaps even simple respiration could have allowed rotation at ~20 degrees per second. Hydrostatic and hydrodynamic properties of torticones suggest that rotation and vertical movement potential constrained the behavior of these helically-coiled cephalopods. We interpret that torticone ammonoids, prominent throughout neritic and epeiric seas during the Albian and Cenomanian (Cretaceous), may have used passive spiral motions to feed upon small food items through the water column, and may have had low metabolic demands compared to modern-day coleoids.