Anatomy and size of Megateuthis, the largest belemnite.

Belemnite rostra are very abundant in Mesozoic marine deposits in many regions. Despite this abundance, soft-tissue specimens of belemnites informing about anatomy and proportions of these coleoid cephalopods are extremely rare and limited to a few moderately large genera like Passaloteuthis and Hibolithes. For all other genera, we can make inferences on their body proportions and body as well as mantle length by extrapolating from complete material. We collected data of the proportions of the hard parts of some Jurassic belemnites in order to learn about shared characteristics in their gross anatomy. This knowledge is then applied to the Bajocian genus Megateuthis, which is the largest known belemnite genus worldwide. Our results provide simple ratios that can be used to estimate belemnite body size, where only the rostrum is known.

'Arm brains' (axial nerves) of Jurassic coleoids and the evolution of coleoid neuroanatomy.

Although patchy, the fossil record of coleoids bears a wealth of information on their soft part anatomy. Here, we describe remains of the axial nerve cord from both decabrachian (Acanthoteuthis, Belemnotheutis, Chondroteuthis) and octobrachian (Plesioteuthis, Proteroctopus, Vampyronassa) coleoids from the Jurassic. We discuss some hypotheses reflecting on possible evolutionary drivers behind the neuroanatomical differentiation of the coleoid arm crown. We also propose some hypotheses on potential links between habitat depth, mode of life and the evolution of the Coleoidea. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00285-3.

Biological light-weight materials: The endoskeletons of cephalopod mollusks.

Structural biological hard tissues fulfill diverse tasks: protection, defence, locomotion, structural support, reinforcement, buoyancy. The cephalopod mollusk Spirula spirula has a planspiral, endogastrically coiled, chambered, endoskeleton consisting of the main elements: shell-wall, septum, adapical-ridge, siphuncular-tube. The cephalopod mollusk Sepia officinalis has an oval, flattened, layered-cellular endoskeleton, formed of the main elements: dorsal-shield, wall/pillar, septum, siphuncular-zone. Both endoskeletons are light-weight buoyancy devices that enable transit through marine environments: vertical (S. spirula), horizontal (S. officinalis). Each skeletal element of the phragmocones has a specific morphology, component structure and organization. The conjunction of the different structural and compositional characteristics renders the evolved nature of the endoskeletons and facilitates for Spirula frequent migration from deep to shallow water and for Sepia coverage over large horizontal distances, without damage of the buoyancy device. Based on Electron-Backscatter-Diffraction (EBSD) measurements and TEM, FE-SEM, laser-confocal-microscopy imaging we highlight for each skeletal element of the endoskeleton its specific mineral/biopolymer hybrid nature and constituent arrangement. We demonstrate that a variety of crystal morphologies and biopolymer assemblies are needed for enabling the endoskeleton to act as a buoyancy device. We show that all organic components of the endoskeletons have the structure of cholesteric-liquid-crystals and indicate which feature of the skeletal element yields the necessary mechanical property to enable the endoskeleton to fulfill its function. We juxtapose structural, microstructural, texture characteristics and benefits of coiled and planar endoskeletons and discuss how morphometry tunes structural biomaterial function. Both mollusks use their endoskeleton for buoyancy regulation, live and move, however, in distinct marine environments.

Cephalopod palaeobiology: evolution and life history of the most intelligent invertebrates: Honoring Sigurd von Boletzky and his contributions to cephalopod research.

Sigurd von Boletzky was a cephalopod researcher who was world-renowned for his enthusiasm for his field of research, for his friendly and calm personality, and, of course, his publications. He dedicated most of his life as active researcher on the development, biology and evolution of coleoids. Nevertheless, he was always curious to learn about other cephalopods as well. Sigurd passed away in Switzerland on September 28th 2020. We dedicate this text and volume to his memory.

Evolutionary development of the cephalopod arm armature: a review.

The cephalopod arm armature is certainly one of the most important morphological innovations responsible for the evolutionary success of the Cephalopoda. New palaeontological discoveries in the recent past afford to review and reassess origin and homology of suckers, sucker rings, hooks, and cirri. Since a priori character state reconstructions are still ambiguous, we suggest and discuss three different evolutionary scenarios. Each of them is based on the following assumptions: (1) Neocoleoidea uniting extant Decabrachia and Octobrachia is monophyletic (= proostracum-bearing coleoids); (2) extinct Belemnitida and Diplobelida are stem decabrachians; (3) proostracum-less coleoids (Hematitida, Donovaniconida, Aulacoceratida) represent stem-neocoleoids; (4) Ammonoidea and Bactritoidea are stem coleoids. We consider a scenario where belemnoid hooks derived from primitive suckers as well-supported. Regarding belemnoid hooks and suckers as homologues implies that belemnoid, oegopsid, and probably ammonoid arm hooks arose through parallel evolution. Our conclusions challenge the widespread opinion, whereupon belemnoid hooks evolved de novo, and instead support earlier ideas formulated by Sigurd von Boletzky.

Fossilized leftover falls as sources of palaeoecological data: a 'pabulite' comprising a crustacean, a belemnite and a vertebrate from the Early Jurassic Posidonia Shale.

Especially in Lagerstätten with exceptionally preserved fossils, we can sometimes recognize fossilized remains of meals of animals. We suggest the term leftover fall for the event and the term pabulite for the fossilized meal when it never entered the digestive tract (difference to regurgitalites). Usually, pabulites are incomplete organismal remains and show traces of the predation. Pabulites have a great potential to inform about predation as well as anatomical detail, which is invisible otherwise. Here, we document a pabulite comprising the belemnite Passaloteuthis laevigata from the Toarcian of the Holzmaden region. Most of its soft parts are missing while the arm crown is one of the best preserved that is known. Its arms embrace an exuvia of a crustacean. We suggest that the belemnite represents the remnant of the food of a predatory fish such as the shark Hybodus.

Preservation of nautilid soft parts inside and outside the conch interpreted as central nervous system, eyes, and renal concrements from the Lebanese Cenomanian.

Nautilid, coleoid and ammonite cephalopods preserving jaws and soft tissue remains are moderately common in the extremely fossiliferous Konservat-Lagerstätte of the Hadjoula, Haqel and Sahel Aalma region, Lebanon. We assume that hundreds of cephalopod fossils from this region with soft-tissues lie in collections worldwide. Here, we describe two specimens of Syrionautilus libanoticus (Cymatoceratidae, Nautilida, Cephalopoda) from the Cenomanian of Hadjoula. Both specimens preserve soft parts, but only one shows an imprint of the conch. The specimen without conch displays a lot of anatomical detail. We homologise the fossilised structures as remains of the digestive tract, the central nervous system, the eyes, and the mantle. Small phosphatic structures in the middle of the body chamber of the specimen with conch are tentatively interpreted as renal concrements (uroliths). The absence of any trace of arms and the hood of the specimen lacking its conch is tentatively interpreted as an indication that this is another leftover fall (pabulite), where a predator lost parts of its prey. Other interpretations such as incomplete scavenging are also conceivable.

Taphonomic patterns mimic biologic structures: diagenetic Liesegang rings in Mesozoic coleoids and coprolites.

Because of physiology of coleoids, their fossils preserve soft-tissue-remains more often than other cephalopods. Sometimes, the phosphatized soft-tissues, particularly parts of the muscular mantle, display dark circular patterns. Here, we showcase that these patterns, here documented for fossil coleoids from the Jurassic of Germany and the Cretaceous of Lebanon, superficially resemble chromatophores (which enable living coleoids to alter their coloration). We examined and chemically analyzed the circular structures in these specimens, describe them, and discuss their genesis. Based on their structure and color, we visually differentiate between three types of circles. By comparison with similar structures, we suggest that these structures are not biogenic but Liesegang rings, which formed due to reaction-diffusion processes very soon after death.

Recent advances in heteromorph ammonoid palaeobiology.

Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non-planispiral coiling. Such aberrant forms appeared convergently four times within this extinct group of cephalopods. Since Wiedmann's seminal paper in this journal, the palaeobiology of heteromorphs has advanced substantially. Combining direct evidence from their fossil record, indirect insights from phylogenetic bracketing, and physical as well as virtual models, we reach an improved understanding of heteromorph ammonoid palaeobiology. Their anatomy, buoyancy, locomotion, predators, diet, palaeoecology, and extinction are discussed. Based on phylogenetic bracketing with nautiloids and coleoids, heteromorphs like other ammonoids had 10 arms, a well-developed brain, lens eyes, a buccal mass with a radula and a smaller upper as well as a larger lower jaw, and ammonia in their soft tissue. Heteromorphs likely lacked arm suckers, hooks, tentacles, a hood, and an ink sac. All Cretaceous heteromorphs share an aptychus-type lower jaw with a lamellar calcitic covering. Differences in radular tooth morphology and size in heteromorphs suggest a microphagous diet. Stomach contents of heteromorphs comprise planktic crustaceans, gastropods, and crinoids, suggesting a zooplanktic diet. Forms with a U-shaped body chamber (ancylocone) are regarded as suspension feeders, whereas orthoconic forms additionally might have consumed benthic prey. Heteromorphs could achieve near-neutral buoyancy regardless of conch shape or ontogeny. Orthoconic heteromorphs likely had a vertical orientation, whereas ancylocone heteromorphs had a near-horizontal aperture pointing upwards. Heteromorphs with a U-shaped body chamber are more stable hydrodynamically than modern Nautilus and were unable substantially to modify their orientation by active locomotion, i.e. they had no or limited access to benthic prey at adulthood. Pathologies reported for heteromorphs were likely inflicted by crustaceans, fish, marine reptiles, and other cephalopods. Pathologies on Ptychoceras corroborates an external shell and rejects the endocochleate hypothesis. Devonian, Triassic, and Jurassic heteromorphs had a preference for deep-subtidal to offshore facies but are rare in shallow-subtidal, slope, and bathyal facies. Early Cretaceous heteromorphs preferred deep-subtidal to bathyal facies. Late Cretaceous heteromorphs are common in shallow-subtidal to offshore facies. Oxygen isotope data suggest rapid growth and a demersal habitat for adult Discoscaphites and Baculites. A benthic embryonic stage, planktic hatchlings, and a habitat change after one whorl is proposed for Hoploscaphites. Carbon isotope data indicate that some Baculites lived throughout their lives at cold seeps. Adaptation to a planktic life habit potentially drove selection towards smaller hatchlings, implying high fecundity and an ecological role of the hatchlings as micro- and mesoplankton. The Chicxulub impact at the Cretaceous/Paleogene (K/Pg) boundary 66 million years ago is the likely trigger for the extinction of ammonoids. Ammonoids likely persisted after this event for 40-500 thousand years and are exclusively represented by heteromorphs. The ammonoid extinction is linked to their small hatchling sizes, planktotrophic diets, and higher metabolic rates than in nautilids, which survived the K/Pg mass extinction event.

Jaws of a large belemnite and an ammonite from the Aalenian (Middle Jurassic) of Switzerland.

Although belemnite rostra can be quite abundant in Jurassic and Cretaceous strata, the record of belemnite jaws was limited to a few specimens from Germany and Russia. Here, we describe and figure three cephalopod jaws from the Middle Jurassic Opalinus Clay of northern Switzerland. Although flattened, the carbonaceous fossils display enough morphological information to rule out an ammonoid, nautiloid or octobrachian origin of the two larger jaws. Their similarities to belemnite jaws from Germany and Russia conforms with our interpretation of these specimens as belemnite jaws. Based on their rather large size, we tentatively assign these two jaws to the megateuthidid Acrocoelites conoideus. The third jaw is a rather small upper jaw of an ammonoid. Since Leioceras opalinum is by far the most common ammonite in this unit in northern Switzerland, we tentatively suggest that the upper jaw belongs to this species.

The shell matrix and microstructure of the Ram's Horn squid: Molecular and structural characterization.

Molluscs are one of the most diversified phyla among metazoans. Most of them produce an external calcified shell, resulting from the secretory activity of a specialized epithelium of the calcifying mantle. This biomineralization process is controlled by a set of extracellular macromolecules, the organic matrix. In spite of several studies, these components are mainly known for bivalves and gastropods. In the present study, we investigated the physical and biochemical properties of the internal planispiral shell of the Ram's Horn squid Spirula spirula. Scanning Electron Microscope investigations of the shell reveal a complex microstructural organization. The saccharides constitute a quantitatively important moiety of the matrix, as shown by Fourier-transform infrared and solid-state nuclear magnetic resonance spectroscopies. NMR identified ß-chitin and additional polysaccharides for a total amount of 80% of the insoluble fraction. Proteomics was applied to both soluble and insoluble matrices and in silico searches were performed, first on heterologous metazoans models, and secondly on an unpublished transcriptome of Spirula spirula. In the first case, several peptides were identified, some of them matching with tyrosinase, chitinase 2, protease inhibitor, or immunoglobulin. In the second case, 39 hits were obtained, including transferrin, a serine protease inhibitor, matrilin, or different histones. The very few similarities with known molluscan shell matrix proteins suggest that Spirula spirula uses a unique set of shell matrix proteins for constructing its internal shell. The absence of similarity with closely related cephalopods demonstrates that there is no obvious phylogenetic signal in the cephalopod skeletal matrix.

Chamber volume development, metabolic rates, and selective extinction in cephalopods.

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 palaeobiology of belemnites - foundation for the interpretation of rostrum geochemistry.

Belemnites are an extinct group of Mesozoic coleoid cephalopods with a fossil record ranging from the early Late Triassic [about 240 million years ago (Mya)] to the Cretaceous/Palaeogene boundary (65 Mya). Belemnites were widely distributed, highly abundant and diverse, and an important component of Mesozoic marine food webs. Their internal shells, specifically their low-Mg calcite rostra, have been used as palaeoenvironmental carbonate archives for the last 70 years. This is primarily due to the assumption that the rostrum calcite formed in equilibrium with the oxygen isotope composition of ambient sea water. Of prime importance for the reliable interpretation of isotope data derived from these biogenic carbonates is a robust reconstruction of the palaeobiology of their producers. Here we provide a critical assessment of published reconstructions of belemnite soft-body organization and their lifestyle and habitats. Different lines of evidence, including sedimentological, geochemical, morphological, and biomechanical data, point towards an outer shelf habitat of belemnites, for some taxa also including the littoral area. Belemnite habitat temperatures, oxygen content, salinities, and life span are constrained based on observations of the ecology and life history of modern coleoids. Belemnite habitat depth might have been largely controlled by food and temperature, with a temperature optimum between 10°C and 30°C. The distribution of modern coleoids is for most species restricted to well-oxygenated water masses and a salinity between 27 and 37 psu. The trophic position of belemnites as both predators and prey is documented by unique fossil finds of stomach contents and soft tissue preservation, such as jaws, hooks, and ink sacs. Belemnites were medium-sized predators in the epipelagic zone (not deeper than ∼200 m) hunting for crustaceans, other cephalopods, and fishes. Taxa with elongated rostra probably were fast and highly manoeuvrable swimmers. Forms with conical rostra represent slow but highly manoeuvrable swimmers, and forms with depressed rostra likely had a bottom-related life habit. Predators of adult belemnites were sharks, bony fishes, and marine reptiles. Belemnites, like most of the modern coleoids, were relatively short lived, most likely living only for 1-2 years. Understanding the biomineralization of belemnite rostra is highly relevant for an improved interpretation of their geochemistry. Here we confirm that belemnite rostra are composed of low Mg-calcite fibres, but they do not contain distinct types of laminae. These fibres are composed of two distinct calcite phases. One phase is a filigree network of tetrahedral organic-rich calcite and the second phase is represented by organic-poor calcite.

Anatomy and evolution of the first Coleoidea in the Carboniferous.

Coleoidea (squids and octopuses) comprise all crown group cephalopods except the Nautilida. Coleoids are characterized by internal shell (endocochleate), ink sac and arm hooks, while nautilids lack an ink sac, arm hooks, suckers, and have an external conch (ectocochleate). Differentiating between straight conical conchs (orthocones) of Palaeozoic Coleoidea and other ectocochleates is only possible when rostrum (shell covering the chambered phragmocone) and body chamber are preserved. Here, we provide information on how this internalization might have evolved. We re-examined one of the oldest coleoids, Gordoniconus beargulchensis from the Early Carboniferous of the Bear Gulch Fossil-Lagerstätte (Montana) by synchrotron, various lights and Reflectance Transformation Imaging (RTI). This revealed previously unappreciated anatomical details, on which we base evolutionary scenarios of how the internalization and other evolutionary steps in early coleoid evolution proceeded. We suggest that conch internalization happened rather suddenly including early growth stages while the ink sac evolved slightly later.

High voltage electrochemical exfoliation of graphite for high-yield graphene production.

We demonstrate a highly efficient, single-step, cathodic exfoliation process of graphite to produce single- to few-layer graphene with a yield of over 70% from natural graphite flakes. By employing boron-doped diamond electrodes high potentials up to -60 V can be applied which was found to greatly increase the yield. The produced graphene flakes are partially hydrogenated during the electrochemical treatment likely aiding in their exfoliation. The resulting flakes have a large lateral size with up to 50 µm diameter. Due to the reversibility of the hydrogenation by thermal treatment the graphene flakes possess a low defect density as judged by the Raman D/G ratio yielding highly conductive films with sheet resistances of 100 to 3200 Ω â–¡-1 at 10 to 70% transparency.

Graphene as an active virtually massless top electrode for RF solidly mounted bulk acoustic wave (SMR-BAW) resonators.

Mechanical and electrical losses induced by an electrode material greatly influence the performance of bulk acoustic wave (BAW) resonators. Graphene as a conducting and virtually massless 2D material is a suitable candidate as an alternative electrode material for BAW resonators which reduces electrode induced mechanical losses. In this publication we show that graphene acts as an active top electrode for solidly mounted BAW resonators (BAW-SMR) at 2.1 GHz resonance frequency. Due to a strong decrease of mass loading and its remarkable electronic properties, graphene demonstrates its ability as an ultrathin conductive layer. In our experiments we used an optimized graphene wet transfer on aluminum nitride-based solidly mounted resonator devices. We achieved more than a triplication of the resonator's quality factor Q and a resonance frequency close to an 'unloaded' resonator without metallization. Our results reveal the direct influence of both, the graphene quality and the graphene contacting via metal structures, on the performance characteristic of a BAW resonator. These findings clearly show the potential of graphene in minimizing mechanical losses due to its virtually massless character. Moreover, they highlight the advantages of graphene and other 2D conductive materials for alternative electrodes in electroacoustic resonators for radio frequency applications.

Wettability Investigations and Wet Transfer Enhancement of Large-Area CVD-Graphene on Aluminum Nitride.

The two-dimensional and virtually massless character of graphene attracts great interest for radio frequency devices, such as surface and bulk acoustic wave resonators. Due to its good electric conductivity, graphene might be an alternative as a virtually massless electrode by improving resonator performance regarding mass-loading effects. We report on an optimization of the commonly used wet transfer technique for large-area graphene, grown via chemical vapor deposition, onto aluminum nitride (AlN), which is mainly used as an active, piezoelectric material for acoustic devices. Today, graphene wet transfer is well-engineered for silicon dioxide (SiO2). Investigations on AlN substrates reveal highly different surface properties compared to SiO2 regarding wettability, which strongly influences the quality of transferred graphene monolayers. Both physical and chemical effects of a plasma treatment of AlN surfaces change wettability and avoid large-scale cracks in the transferred graphene sheet during desiccation. Spatially-resolved Raman spectroscopy reveals a strong strain and doping dependence on AlN plasma pretreatments correlating with the electrical conductivity of graphene. In our work, we achieved transferred crack-free large-area (40 × 40 mm²) graphene monolayers with sheet resistances down to 350 Ω/sq. These achievements make graphene more powerful as an eco-friendly and cheaper replacement for conventional electrode materials used in radio frequency resonator devices.

Microtomography of the Baltic amber tick Ixodes succineus reveals affinities with the modern Asian disease vector Ixodes ovatus.

BACKGROUND: Fossil ticks are extremely rare and Ixodes succineus Weidner, 1964 from Eocene (ca. 44-49 Ma) Baltic amber is one of the oldest examples of a living hard tick genus (Ixodida: Ixodidae). Previous work suggested it was most closely related to the modern and widespread European sheep tick Ixodes ricinus (Linneaus, 1758). RESULTS: Restudy using phase contrast synchrotron x-ray tomography yielded images of exceptional quality. These confirm the fossil's referral to Ixodes Latreille, 1795, but the characters resolved here suggest instead affinities with the Asian subgenus Partipalpiger Hoogstraal et al., 1973 and its single living (and medically significant) species Ixodes ovatus Neumann, 1899. We redescribe the amber fossil here as Ixodes (Partipalpiger) succineus. CONCLUSIONS: Our data suggest that Ixodes ricinus is unlikely to be directly derived from Weidner's amber species, but instead reveals that the Partipalpiger lineage was originally more widely distributed across the northern hemisphere. The closeness of Ixodes (P.) succineus to a living vector of a wide range of pathogens offers the potential to correlate its spatial and temporal position (northern Europe, nearly 50 million years ago) with the estimated origination dates of various tick-borne diseases.