@article{HoffmannLemanisFalkenbergetal., author = {Hoffmann, Ren{\´e} and Lemanis, Robert and Falkenberg, Janina and Schneider, Steffen and Wesendonk, Hendrik and Zachow, Stefan}, title = {Integrating 2D and 3D shell morphology to disentangle the palaeobiology of ammonoids: A virtual approach}, series = {Palaeontology}, volume = {61}, journal = {Palaeontology}, number = {1}, doi = {10.1111/pala.12328}, pages = {89 -- 104}, abstract = {We demonstrate, based on data derived from computed tomography, that integrating 2D and 3D morphological data of ammonoid shells represent an important new approach to disentangle the palaeobiology of ammonoids. Characterization of ammonite morphology has long been constrained to 2D data and only a few studies collect ontogenetic data in 180° steps. We combine this traditional approach with 3D data collected from high-resolution nano-computed tomography. For this approach, ontogenetic morphological data of the hollow shell of a juvenile ammonite Kosmoceras (Jurassic, Callovian) is collected. 2D data is collected in 10° steps and show significant changes in shell morphology. Preserved hollow spines show multiple mineralized membranes never reported before, representing temporal changes of the ammonoid mantle tissue. 3D data show that chamber volumes do not always increase exponentially, as was generally assumed, but may represent a proxy for life events, e.g. stress phases. Furthermore, chamber volume cannot be simply derived from septal spacing in forms comparable to Kosmoceras. Vogel numbers, a 3D parameter for chamber shape, of Kosmoceras are similar to other ammonoids (Arnsbergites, Amauroceras) and modern cephalopods (Nautilus, Spirula). Two methods to virtually document the suture line ontogeny, used to document phylogenetic relationships of larger taxonomic entities, were applied for the first time and seem a promising alternative to hand drawings. The curvature of the chamber surfaces increases during ontogeny due to increasing strength of ornamentation and septal complexity. As increasing curvature may allow for faster handling of cameral liquid, it could compensate for decreasing SA/V ratios through ontogeny.}, language = {en} } @article{LemanisKornZachowetal., author = {Lemanis, Robert and Korn, Dieter and Zachow, Stefan and Rybacki, Erik and Hoffmann, Ren{\´e}}, title = {The Evolution and Development of Cephalopod Chambers and Their Shape}, series = {PLOS ONE}, volume = {11}, journal = {PLOS ONE}, number = {3}, doi = {10.1371/journal.pone.0151404}, abstract = {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.}, language = {en} } @article{LemanisZachowHoffmann, author = {Lemanis, Robert and Zachow, Stefan and Hoffmann, Ren{\´e}}, title = {Comparative cephalopod shell strength and the role of septum morphology on stress distribution}, series = {PeerJ}, volume = {4}, journal = {PeerJ}, doi = {10.7717/peerj.2434}, pages = {e2434}, 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.}, language = {en} } @article{HoffmannSchultzSchellhornetal., author = {Hoffmann, Ren{\´e} and Schultz, Julia A. and Schellhorn, Rico and Rybacki, Erik and Keupp, Helmut and Gerden, S. R. and Lemanis, Robert and Zachow, Stefan}, title = {Non-invasive imaging methods applied to neo- and paleontological cephalopod research}, series = {Biogeosciences}, volume = {11}, journal = {Biogeosciences}, number = {10}, doi = {10.5194/bg-11-2721-2014}, pages = {2721 -- 2739}, 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.}, language = {en} } @article{LemanisZachowFusseisetal., author = {Lemanis, Robert and Zachow, Stefan and Fusseis, Florian and Hoffmann, Ren{\´e}}, title = {A new approach using high-resolution computed tomography to test the buoyant properties of chambered cephalopod shells}, series = {Paleobiology}, volume = {41}, journal = {Paleobiology}, number = {2}, publisher = {Cambridge University Press}, address = {Cambridge}, doi = {10.1017/pab.2014.17}, pages = {313 -- 329}, abstract = {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 on Nautilus pompilius and is now extended to the extant deep-sea squid Spirula spirula and the Jurassic ammonite Cadoceras sp. hatchling. Cadoceras is 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 of Cadoceras supports 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.}, language = {en} } @article{HoffmannLemanisWulffetal., author = {Hoffmann, Rene and Lemanis, Robert and Wulff, Lena and Zachow, Stefan and Lukeneder, Alexander and Klug, Christian and Keupp, Helmut}, title = {Traumatic events in the life of the deep-sea cephalopod mollusc, the coleoid Spirula spirula}, series = {ScienceDirect: Deep Sea Research Part I - Oceanographic Research}, volume = {142}, journal = {ScienceDirect: Deep Sea Research Part I - Oceanographic Research}, number = {12}, doi = {10.1016/j.dsr.2018.10.007}, pages = {127 -- 144}, abstract = {Here, we report on different types of shell pathologies of the enigmatic deep-sea (mesopelagic) cephalopod Spirula spirula. For the first time, we apply non-invasive imaging methods to: document trauma-induced changes in shell shapes, reconstruct the different causes and effects of these pathologies, unravel the etiology, and attempt to quantify the efficiency of the buoyancy apparatus. We have analysed 2D and 3D shell parameters from eleven shells collected as beach findings from the Canary Islands (Gran Canaria and Fuerteventura), West-Australia, and the Maldives. All shells were scanned with a nanotom-m computer tomograph. Seven shells were likely injured by predator attacks: fishes, cephalopods or crustaceans, one specimen was infested by an endoparasite (potentially Digenea) and one shell shows signs of inflammation and one shell shows large fluctuations of chamber volumes without any signs of pathology. These fluctuations are potential indicators of a stressed environment. Pathological shells represent the most deviant morphologies of a single species and can therefore be regarded as morphological end-members. The changes in the shell volume / chamber volume ratio were assessed in order to evaluate the functional tolerance of the buoyancy apparatus showing that these had little effect.}, language = {en} } @misc{HoffmannSchultzSchellhornetal., author = {Hoffmann, Ren{\´e} and Schultz, Julia A. and Schellhorn, Rico and Rybacki, Erik and Keupp, Helmut and Lemanis, Robert and Zachow, Stefan}, title = {Non-invasive imaging methods applied to neo- and paleo-ontological cephalopod research}, issn = {1438-0064}, doi = {10.5194/bg-11-2721-2014}, url = {http://nbn-resolving.de/urn:nbn:de:0297-zib-50300}, 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.}, language = {en} }