Spirula spirula

Abstract: The evolution of complexly folded septa in ammonoids has long been a controversial topic. Explanations of the function of these folded septa can be divided into physiological and mechanical hypotheses with the mechanical functions tending to find widespread support. The complexity of the cephalopod shell has made it difficult to directly test the mechanical properties of these structures without oversimplification of the septal morphology or extraction of a small sub-domain. However, the power of modern finite element analysis now permits direct testing of mechanical hypothesis on complete, empirical models of the shells taken from computed tomographic data. Here we compare, for the first time using empirical models, the capability of the shells of extant Nautilus pompilius, Spirula spirula, and the extinct ammonite Cadoceras sp. to withstand hydrostatic pressure and point loads. Results show hydrostatic pressure imparts highest stress on the final septum with the rest of the shell showing minimal compression. S. spirula shows the lowest stress under hydrostatic pressure while N. pompilius shows the highest stress. Cadoceras sp. shows the development of high stress along the attachment of the septal saddles with the shell wall. Stress due to point loads decreases when the point force is directed along the suture as opposed to the unsupported chamber wall. Cadoceras sp. shows the greatest decrease in stress between the point loads compared to all other models. Greater amplitude of septal flutes corresponds with greater stress due to hydrostatic pressure; however, greater amplitude decreases the stress magnitude of point loads directed along the suture. In our models, sutural complexity does not predict greater resistance to hydrostatic pressure but it does seem to increase resistance to point loads, such as would be from predators. This result permits discussion of palaeoecological reconstructions on the basis of septal morphology. We further suggest that the ratio used to characterize septal morphology in the septal strength index and in calculations of tensile strength of nacre are likely insufficient. A better understanding of the material properties of cephalopod nacre may allow the estimation of maximum depth limits of shelled cephalopods through finite element analysis.

Abstract: The ram’s horn squid, Spirula spirula (Spirulida, Coleoidea), inhabits subsurface waters of the tropical and subtropical oceans. Because of the presence of an internal chambered shell in this species, it has frequently been used as a model species to investigate the paleobiology of fossil coleoids. However, the feeding and dietary habits of S. spirula in the nature are poorly known. In this study, we applied a new method (amino acid nitrogen isotopic analysis) to estimate the trophic position of S. spirula specimens captured off Suriname, as well as three cuttlefish Sepia species (Sepia officinalis, S. latimanus, and S. esculenta), with a calcified chambered shell from the shallower water. The trophic position of S. spirula was estimated to be 2.5–2.8, which was significantly lower than that for the three Sepia species (3.4–3.6). The results and available data on the gastric contents of S. spirula suggest that it feeds mainly on detritus and zooplankton, including crustaceans, from the overlying water column. The method used in this study can potentially be applied to the estimation of the trophic position of the fossil cephalopods having calcified chambered shells.