René Hoffmann

Ammonoid Intraspecific Variability

Two main types of intraspecific variation can be distinguished in ammonoids, which are not mutually exclusive: continuous and discontinuous variation. Although many authors acknowledge or implicitly assume a large intraspecific variability is possible in shell shape, ornamentation and suture line, it has only been rarely studied quantitatively. Several potential biases need to be taken into account when studying intraspecific variation of fossil populations including paleoecological, taphonomic and collection biases. Intraspecific variation might be controlled both by genetic and environmental parameters, although both are difficult to separate in fossil samples. In ammonoids, a large part of intraspecific variation in morphology and size has been attributed to differences in growth rates and development. Taking intraspecific variation properly into account is not only of prime importance for taxonomy, but also for studies on biostratigraphy, paleobiogeography, ecology, paleobiology and evolution of ammonoids.

A new approach using high-resolution computed tomography to test the buoyant properties of chambered cephalopod shells

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.

Comparative cephalopod shell strength and the role of septum morphology on stress distribution

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.

Ammonoid Septa and Sutures

Ranging from simple to complex, the folded septa of the phragmocone characterize the ammonoid clade and contribute much to their aesthetic appearance. However, many contradicting opinions and models on septum formation and as explanation for the evolution and function of sutural complexity and septal frilling have been published. We review the main hypotheses, namely the Viscous Fingering Model, the Tie-Point Model and the application of the Reaction Diffusion Model to the morphogenesis of ammonoid septa. In that context, we discuss contradictions and present a compound model including a revised chamber formation cycle. Additionally, the highly complex septa of lytoceratids are discussed in greater details with respect to septum formation and the evolutionary meaning of suture lines and septum morphology.

The Evolution and Development of Cephalopod Chambers and Their Shape

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.

Non-invasive imaging techniques combined with morphometry: a case study from Spirula

Spirula spirula is a unique deep-sea squid with unknown taxonomic status. Precise description of shell morphology may help to decide whether the genus contains one or more species. Here, a straight forward description of ontogenetic changes of shell parameters is presented for a single shell of Spirula spirula. Using micro-computed tomography, surface and volumetric data, e.g., chamber volumes and surface areas, as well as siphuncle volumes and surface areas were collected and used for the description. Advantage of the method, combining non-invasive imaging techniques with classical morphometry, is discussed.

Two New Ammonoid Genera of the Subfamily Gabbioceratinae from the Upper Albian (Lower Cretaceous) of Hokkaido, Japan

Abstract. Two ammonoids of the subfamily Gabbioceratinae, Obataceras manjiense gen. et sp. nov. and Tanabeceras pombetsense gen. et sp. nov., are described from the Upper Albian of Hokkaido, Japan. The Gabbioceratinae evolved and radiated mainly in the Mediterranean area during Late Aptian to Middle Albian times, but thereafter disappeared from the region. The occurrences of Gabbioceratinae in Hokkaido suggest that their geographical distribution was extended to the Northwest Pacific before the Late Albian, where they then flourished from Late Albian to Cenomanian time.

Grasping the shape of belemnoid arm hooks : a quantitative approach

Abstract. Chitinous arm hooks (onychites) of belemnoid coleoid cephalopods are widely distributed in Mesozoic sediments. Due to their relative abundance and variable morphology compared with the single, bullet-shaped, belemnite rostrum, arm hooks came into the focus of micropaleontologists as a promising index fossil group for the Jurassic—Cretaceous rock record and have been the target of functional, ecological, and phylogenetic interpretations in the past. Based on three well-preserved arm crowns of the Toarcian diplobelid Chondroteuthis wunnenbergi, we analyzed the shape of a total of 87micro-hooks. The arm crown of Chondroteuthis is unique in having uniserial rather than biserial hooks. The first application of elliptic Fourier shape analysis to the arm weapons of belemnoid coleoids allows for the distinction of four micro-hook morphotypes and the quantification of shape variation within these morphotypes. Based on the best-preserved arm crown, we reconstructed the distribution of morphotypes within the arm crown and along a single arm. Our quantitative data support former observations that smaller hookswere found close to the mouth and at the most distal arm parts, while the largest hooks were found in the central part of the arm crown. Furthermore, we found a distinct arm differentiation, as not every arm was equipped with the same hook morphotype. Here, we report the functional specialization of the belemnoid arm crown for the first time and speculate about the potential function of the four morphotypes based on comparisons with modern cephalopods. Our analyses suggest a highly adapted functional morphology and intra-individual distribution of belemnoid hooks serving distinct purposes mainly during prey capture.

Form and Formation of Flares and Parabolae Based on New Observations of the Internal Shell Structure in Lytoceratid and Perisphinctid Ammonoids

The ultrastructure of pristine shells of Jurassic and Cretaceous lytoceratid and perisphinctid ammonoids indicates that flares and parabolae represent homologous structures. Both mark an interruption of shell growth. We dismiss earlier interpretations of parabolae as actual aperture, relics of resorbed apophyses or superstructure of the musculature associated to a semi-internal shell. Instead we propose an episodic growth model including several growth stops at the aperture during the formation of a frill-like aperture for parabolae and flares. Such an aperture is composed of the outer prismatic layer, the nacreous layer and an apertural prismatic coating. Here, we observed the apertural prismatic coating for the first time as an integral part of flares and parabolae. The apertural prismatic coating covers only the inner surface of the frill and was secreted by a permanent mantle cover indicating a prolonged period without the production of new shell material. Parabolae differ from flares by their general shape and the presence of ventro-lateral parabolic notches and nodes. The notches were formed by folding of the frill and had the potential to form semi-open spines. The corresponding parabolic nodes are caused by an outward swelling of the shell-secreting mantle tissue producing new shell material at the position of the folding. New shell material that belongs to the conch tube is attached to the base of flares and parabolae after withdrawal of the mantle edge representing the continuation of shell growth. Usually, the frilled aperture associated with flares and parabolae were removed during lifetime. This study reports on flares in Argonauticeras for the first time. In this genus they are typically associated with varices.

The Correct Taxon Name, Authorship, and Publication Date of Extant Ten-Armed Coleoids

ABSTRACT A variety of conflicting names with different authorship is available and has been repeatedly cited for living ten-armed coleoid cephalopods. Here, I review the primary literature and show the correct name, authorship, and date for ten-armed coleoids.