The chambered shell of modern cephalopods functions as a buoyancy apparatus, allowing the animal to enter the water column without expending a large amount of energy to overcome its own weight. Indeed, the chambered shell is largely considered a key adaptation that allowed the earliest cephalopods to leave the ocean floor and enter the water column. It has been argued by some, however, that the iconic chambered shell of Paleozoic and Mesozoic ammonoids did not provide a sufficiently buoyant force to compensate for the weight of the entire animal, thus restricting ammonoids to a largely benthic lifestyle reminiscent of some octopods. Here we develop a technique using high-resolution computed tomography to quantify the buoyant properties of chambered shells without reducing the shell to ideal spirals or eliminating inherent biological variability by using mathematical models that characterize past work in this area. This technique has been tested onNautilus pompiliusand is now extended to the extant deep-sea squidSpirula spirulaand the Jurassic ammoniteCadocerassp. hatchling.Cadocerasis found to have possessed near-neutral to positive buoyancy if hatched when the shell possessed between three and five chambers. However, we show that the animal could also overcome degrees of negative buoyancy through swimming, similar to the paralarvae of modern squids. These calculations challenge past inferences of benthic life habits based solely on calculations of negative buoyancy. The calculated buoyancy ofCadocerassupports the possibility of planktonic dispersal of ammonite hatchlings. This information is essential to understanding ammonoid ecology as well as biotic interactions and has implications for the interpretation of geochemical data gained from the isotopic analysis of the shell.

Molluscan shells are a classic model system to study formation–structure–function relationships in biological materials and the process of biomineralized tissue morphogenesis. Typically, each shell consists of a number of highly mineralized ultrastructures, each characterized by a specific 3D mineral–organic architecture. Surprisingly, in some cases, despite the lack of a mutual biochemical toolkit for biomineralization or evidence of homology, shells from different independently evolved species contain similar ultrastructural motifs. In the present study, using a recently developed physical framework, which is based on an analogy to the process of directional solidification and simulated by phase-field modeling, we compare the process of ultrastructural morphogenesis of shells from 3 major molluscan classes: A bivalve Unio pictorum, a cephalopod Nautilus pompilius, and a gastropod Haliotis asinina. We demonstrate that the fabrication of these tissues is guided by the organisms by regulating the chemical and physical boundary conditions that control the growth kinetics of the mineral phase. This biomineralization concept is postulated to act as an architectural constraint on the evolution of molluscan shells by defining a morphospace of possible shell ultrastructures that is bounded by the thermodynamics and kinetics of crystal growth.

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. Several non-invasive methods are common practice in natural sciences today. Here we present how they can be applied and contribute to current topics in cephalopod (paleo-) biology. Different methods will be compared in terms of time necessary to acquire the data, amount of data, accuracy/resolution, minimum/maximum size of objects that can be studied, the degree of post-processing needed and availability. The main application of the methods is seen in morphometry and volumetry of cephalopod shells. In particular we present a method for precise buoyancy calculation. Therefore, cephalopod shells were scanned together with different reference bodies, an approach developed in medical sciences. It is necessary to know the volume of the reference bodies, which should have similar absorption properties like the object of interest. Exact volumes can be obtained from surface scanning. Depending on the dimensions of the study object different computed tomography techniques were applied.

The Ammonoidea is a group of extinct cephalopods ideal to study evolution through deep time. The evolution of the planispiral shell and complexly folded septa in ammonoids has been thought to have increased the functional surface area of the chambers permitting enhanced metabolic functions such as: chamber emptying, rate of mineralization and increased growth rates throughout ontogeny. Using nano-computed tomography and synchrotron radiation based micro-computed tomography, we present the first study of ontogenetic changes in surface area to volume ratios in the phragmocone chambers of several phylogenetically distant ammonoids and extant cephalopods. Contrary to the initial hypothesis, ammonoids do not possess a persistently high relative chamber surface area. Instead, the functional surface area of the chambers is higher in earliest ontogeny when compared to Spirula spirula. The higher the functional surface area the quicker the potential emptying rate of the chamber; quicker chamber emptying rates would theoretically permit faster growth. This is supported by the persistently higher siphuncular surface area to chamber volume ratio we collected for the ammonite Amauroceras sp. compared to either S. spirula or nautilids. We demonstrate that the curvature of the surface of the chamber increases with greater septal complexity increasing the potential refilling rates. We further show a unique relationship between ammonoid chamber shape and size that does not exist in S. spirula or nautilids. This view of chamber function also has implications for the evolution of the internal shell of coleoids, relating this event to the decoupling of soft-body growth and shell growth.

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Evidence for a composite organic-inorganic fabric of belemnite rostra: Implications for palaeoceanography and palaeoecology, Sedimentary Geology (2016),

Reconstructing the physiology of extinct organisms is key to understanding mechanisms of selective extinction during biotic crises. Soft tissues of extinct organisms are rarely preserved and, therefore, a proxy for physiological aspects is needed. Here, we examine whether cephalopod conchs yield information about their physiology by assessing how the formation of chambers respond to external stimuli such as environmental changes. We measured chamber volume through ontogeny to detect differences in the pattern of chamber volume development in nautilids, coleoids, and ammonoids. Results reveal that the differences between ontogenetic trajectories of these cephalopods involve the presence or absence of abrupt decreases of chamber volume. Accepting the link between metabolic rate and growth, we assume that this difference is rooted in metabolic rates that differ between cephalopod clades. High metabolic rates combined with small hatching size in ammonoids as opposed to lower metabolic rates and much larger hatchlings in most nautilids may explain the selective extinction of ammonoids as a consequence of low food availability at the end of the cretaceous.The Ammonoidea is a group of ectocochleate cephalopods that were extant for more than 350 million years, during which time they played an essential ecological role in the world's oceans as a result of their high abundance, wide distribution, and great diversity. Although they survived several of the most severe mass extinction events in the course of their evolution 1-3 , they perished at the end of the Cretaceous 4,5 . Despite extensive discussion on the selectivity of the K/Pg extinction, the actual mechanisms that led ammonoids to extinction and allowed nautilids to survive, have not yet been fully revealed, although both intrinsic (e.g., smaller embryonic sizes, larger geographical range, and microphagous feeding 4,6,7 ) and extrinsic factors (e.g., surface ocean acidification and global cooling 8,9 ) have been proposed. Details about intrinsic (anatomical and physiological) aspects such as the muscular system and metabolic rates are difficult to assess in extinct organisms because the soft tissue is rarely fossilized 10 . Thus, we need proxies for biological and physiological aspects to fully reveal the actual kill mechanism of ammonoids at the K/Pg boundary. In fact, such biological and physiological traits are apparently strongly linked to macroecology and macroevolution of organisms. For instance, Strotz et al. 11 discovered a significant difference between basal metabolic rates of extinct and extant taxa. Additionally, Payne et al. 12 demonstrated a new perspective on the evolution of bivalves and brachiopods by calculating their metabolic rates. Reconstructing the biological and physiological traits of extinct ammonoids may, therefore, be a key to understanding selective extinction 13 .Most mollusk conchs contain a wealth of information about their development because the entire life history is recorded within the shell. In ectocochleate cephalopods (ammonoi...