Orthoceratoid and coleoid cephalopods from the Middle Triassic of Switzerland with an updated taxonomic framework for Triassic Orthoceratoidea.

Orthoconic cephalopods are subordinate, but persistent, widespread and regionally abundant components of Triassic marine ecosystems. Here, we describe unpublished specimens from the Anisian (Middle Triassic) Besano Formation at Monte San Giorgio, Switzerland. They can be assigned to two major but unrelated lineages, the Coleoidea and the Orthoceratoidea. The orthoceratoids belong to Trematoceras elegans (Münster, 1841) and occur regularly within the Besano Formation, are uniform in size, and have few available morphological characters. In contrast, coleoids are more diverse and appear to be restricted to shorter intervals. A new coleoid is described as Ticinoteuthis chuchichaeschtli gen. et sp. nov. To better put the orthoceratoids of the Besano Formation into perspective, we also synthesise the current taxonomy of Triassic orthoceratoids on a global scale. The currently used scheme is largely outdated, with very little taxonomic progress in the past 100 years. Despite previous research showing the distinctness of Triassic orthoceratoids from Palaeozoic taxa, they are still commonly labelled as "Orthoceras" or "Michelinoceras", which are confined to the Palaeozoic. We show that Triassic orthoceratoids probably belong to a single lineage, the Trematoceratidae, which can be assigned to the Pseudorthocerida based on the embryonic shell and endosiphuncular deposits. Many Triassic species can probably be assigned to Trematoceras, but there are at least two additional Triassic orthoceratoid genera, Paratrematoceras and Pseudotemperoceras. Finally, we review the palaeobiogeographic and stratigraphic distribution of the group and outline possible future research directions.

Oldest record of Machimosaurini (Thalattosuchia, Teleosauroidea): teeth and scavenging traces from the Middle Jurassic (Bajocian) of Switzerland.

The Jurassic period was a time of major diversification for Mesozoic marine reptiles, including Ichthyosauria, Plesiosauria and thalattosuchian Crocodylomorpha. The latter originated in the Early Jurassic and thrived during the Late Jurassic. Unfortunately, the Middle Jurassic, a crucial time in their evolution, has a poor fossil record. Here, we document the first evidence of macrophagous/durophagous Machimosaurini-tribe teleosauroid thalattosuchians from the late Bajocian (ca 169 Ma) in the form of three robust tooth crowns with conical blunt shapes and anastomosed pattern of thick enamel ridges towards the apex, associated with the skeleton of a large ichthyosaur lacking preserved tooth crowns. The tooth crowns were found on the posterior section of the lower jaw (left angular), a lacrimal and the axis neural arch of the ichthyosaur. In addition, some of the distal sections of the posterior dorsal ribs of the ichthyosaur skeleton exhibit rounded bite marks and some elongated furrows that fit in size and shape with the Machimosaurini teeth. These marks, together with the absence of healing in the rib bone are interpreted here as the indicators of peri- to post-mortem scavenging by a Machimosaurini teleosauroid after the large ichthyosaur carcass settled on the floor of a shallow ocean.

A large new Middle Jurassic ichthyosaur shows the importance of body size evolution in the origin of the Ophthalmosauria.

The Middle Jurassic is an important time period for the evolutionary history of marine reptiles as it represented a transitional phase for many clades. Notably, in ichthyosaurs, many early parvipelvian taxa went extinct. The Middle Jurassic saw the emergence of the derived Ophthalmosauria, ultimately becoming the dominant ichthyosaurian clade by the end of the epoch. Even though this is an important period in the evolutionary history of Ophthalmosauria, our understanding remains limited in terms of morphology and taxonomy due to the scarcity of vertebrate-bearing strata. Here we present a large new ichthyosaur from the Bajocian of Switzerland, represented by an almost complete skull with 3D-preserved bones, the (inter)clavicles and a large portion of the postcranial skeleton. After CT- and surface scanning, we reconstructed the 3D in vivo morphology. Our morphological observations and phylogenetic analyses show that the new taxon named Argovisaurus martafernandezi is nested at the base of the Ophthalmosauria. The holotype and only known specimen of Argovisaurus likely represents an adult individual. Bajocian members of the Ophthalmosauria (Mollesaurus and Argovisaurus) were large-bodied animals, a trait typically associated with the more derived Platypterygiinae. This hints at the importance of a large body size early in ophthalmosaurian evolution.LSID: urn:lsid:zoobank.org:act:C3312628-1544-4B87-BBE3-B12346A30BE3LSID: urn:lsid:zoobank.org:act:23C2BD71-8CF0-4D99-848A-0D631518415B.

Plectronoceratids (Cephalopoda) from the latest Cambrian at Black Mountain, Queensland, reveal complex three-dimensional siphuncle morphology, with major taxonomic implications.

The Plectronoceratida includes the earliest known cephalopod fossils and is thus fundamental to a better understanding of the origin and early evolution of this group of molluscs. The bulk of described material comes from the late Cambrian Fengshan Formation in North China with isolated occurrences in South China, Laurentia, Kazakhstan and Siberia. Knowledge of their morphology and taxonomy is limited in that most specimens were only studied as longitudinal sections, which are prone to misinterpretations due to variations in the plane of section. We describe more than 200 new specimens, which exceeds the entire hitherto published record of plectronoceratids. The material was collected by Mary Wade and colleagues during the 1970s and 1980s, from the lower Ninmaroo Formation at Black Mountain (Mount Unbunmaroo), Queensland, Australia. Despite the collecting effort, diverse notes and early incomplete drafts, Mary Wade never published this material before her death in 2005. The specimens provide novel insights into the three-dimensional morphology of the siphuncle based on abundant material, prompting a general revision of the order Plectronoceratida. We describe Sinoeremoceras marywadeae sp. nov. from numerous, well-preserved specimens, allowing investigation of ontogenetic trajectories and intraspecific variability, which in turn enables improved interpretations of the three-dimensional siphuncle morphology. The siphuncle of S. marywadeae sp. nov. and other plectronoceratids is characterised by highly oblique segments, an elongated middorsal portion of the septal neck (= septal flap) and laterally expanded segments that extend dorsally relative to the septal flap (= siphuncular bulbs). We show that this complex siphuncular structure has caused problems of interpretation because it was studied mainly from longitudinal sections, leading to the impression that there were large differences between specimens and supposed species. We revise the order Protactinoceratida and the families Protactinoceratidae and Balkoceratidae as junior synonyms of the Plectronoceratida and Plectronoceratidae, respectively. We reduce the number of valid genera from eighteen (including one genus formerly classified as an ellesmeroceratid) to three: Palaeoceras Flower, 1954, Plectronoceras Kobayashi, 1935 and Sinoeremoceras Kobayashi, 1933. We accept 10 valid species to which the 68 previously established species may be assigned. Sinoeremoceras contains 8 of the 10 plus the new species. Two species, previously referred to ellesmeroceratid genera, are transferred to Sinoeremoceras. This revised scheme groups plectronoceratids into distinct geographically and stratigraphically separated species, which better reflects biological realities and removes bias caused by preparation techniques. North China remains important containing the highest known diversity and was likely a centre of cephalopod diversification.

The marine conservation deposits of Monte San Giorgio (Switzerland, Italy): the prototype of Triassic black shale Lagerstätten.

Marine conservation deposits ('Konservat-Lagerstätten') are characterized by their mode of fossil preservation, faunal composition and sedimentary facies. Here, we review these characteristics with respect to the famous conservation deposit of the Besano Formation (formerly Grenzbitumenzone; including the Anisian-Ladinian boundary), and the successively younger fossil-bearing units Cava inferiore, Cava superiore, Cassina beds and the Kalkschieferzone of Monte San Giorgio (Switzerland and Italy). We compare these units to a selection of important black shale-type Lagerstätten of the global Phanerozoic plus the Ediacaran in order to detect commonalities in their facies, genesis, and fossil content using principal component and hierarchical cluster analyses. Further, we put the Monte San Giorgio type Fossillagerstätten into the context of other comparable Triassic deposits worldwide based on their fossil content. The results of the principal component and cluster analyses allow a subdivision of the 45 analysed Lagerstätten into four groups, for which we suggest the use of the corresponding pioneering localities: Burgess type for the early Palaeozoic black shales, Monte San Giorgio type for the Triassic black shales, Holzmaden type for the pyrite-rich black shales and Solnhofen type for platy limestones.

Extreme lower jaw elongation in a placoderm reflects high disparity and modularity in early vertebrate evolution.

Jaws are a key vertebrate feature that arose early in our evolution. Placoderms are among the first jawed vertebrates; their fossils yield essential knowledge about the early diversification of gnathostome feeding strategies, diets and modularity. Modularity can be expressed through disproportional lengths of lower and upper jaws as in swordfish or halfbeaks. Alienacanthus malkowskii is an arthrodire from the Famennian of Morocco and Poland, whose most remarkable feature is its lower jaw, which is twice as long as the skull. This is the oldest record of such extreme jaw elongation and modularity in vertebrates. The gnathal plates of Alienacanthus possess sharp, posteriorly recurved teeth that continue anterior of the occlusion in the inferognathals. The dentition suggests a catching and trapping live prey function, and the jaw occlusion is unique among placoderms. This armoured 'fish' expands the morphological and ecological diversity during one of the first radiations of jawed vertebrates with a combination of features so far unrecorded for arthrodires.

'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.

Broad snouted cladoselachian with sensory specialization at the base of modern chondrichthyans.

Throughout the Silurian and Devonian, cartilaginous fish successively evolved their specialized skeletal and dental characteristics, and increasingly refined their sensory systems. The Late Devonian shark taxon Maghriboselache mohamezanei gen. et sp. n. from the eastern Anti-Atlas of Morocco is known from multiple specimens preserving most of its skeletal features, which in some instances are preserved in three dimensions. Key details of the dentition, jaws, and pectoral skeleton are shared with the iconic genus Cladoselache. Phylogenetic analyses place the family Cladoselachidae as the sister group of symmoriiforms and these groups as sister group of the holocephalans. Further phylogenetic results corroborate that the initial evolutionary radiation of crown chondrichthyans occurred within or before the Late Devonian. Remarkably, this new stem holocephalan is equipped with a wide snout and large laterally separated nasal capsules: the earliest known example of this condition in the chondrichthyan and (perhaps) gnathostome record. This suggests sensory specialization approaching that of extant broad-rostrum elasmobranchs and represents a significant addition to increasingly apparent ecomorphological diversity among early chondrichthyans. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00266-6.

A new Meckel's cartilage from the Devonian Hangenberg black shale in Morocco and its position in chondrichthyan jaw morphospace.

Fossil chondrichthyan remains are mostly known from their teeth, scales or fin spines only, whereas their cartilaginous endoskeletons require exceptional preservational conditions to become fossilized. While most cartilaginous remains of Famennian (Late Devonian) chondrichthyans were found in older layers of the eastern Anti-Atlas, such fossils were unknown from the Hangenberg black shale (HBS) and only a few chondrichthyan teeth had been found therein previously. Here, we describe a Meckel's cartilage from the Hangenberg black shale in Morocco, which is the first fossil cartilage from these strata. Since no teeth or other skeletal elements have been found in articulation, we used elliptical Fourier (EFA), principal component (PCA), and hierarchical cluster (HCA) analyses to morphologically compare it with 41 chondrichthyan taxa of different size and age and to evaluate its possible systematic affiliation. PCA and HCA position the new specimen closest to some acanthodian and elasmobranch jaws. Accordingly, a holocephalan origin was excluded. The jaw shape as well as the presence of a polygonal pattern, typical for tessellated calcified cartilage, suggest a ctenacanth origin and we assigned the new HBS Meckel's cartilage to the order Ctenacanthiformes with reservations.

Phosphatized adductor muscle remains in a Cenomanian limid bivalve from Villers-sur-Mer (France).

Soft-tissue preservation in molluscs is generally rare, particularly in bivalves and gastropods. Here, we report a three-dimensionally preserved specimen of the limid Acesta clypeiformis from the Cenomanian of France that shows preservation of organic structures of the adductor muscles. Examination under UV-light revealed likely phosphatisation of organic remains, which was corroborated by EDX-analyses. We suggest that the parts of the adductor muscles that are very close to the attachment are particularly resistant to decay and thus may be preserved even under taphonomic conditions usually not favouring soft-tissue fossilisation.

Is the relative thickness of ammonoid septa influenced by ocean acidification, phylogenetic relationships and palaeogeographic position?

The impact of increasing atmospheric CO2 and the resulting decreasing pH of seawater are in the focus of current environmental research. These factors cause problems for marine calcifiers such as reduced calcification rates and the dissolution of calcareous skeletons. While the impact on recent organisms is well established, little is known about long-term evolutionary consequences. Here, we assessed whether ammonoids reacted to environmental change by changing septal thickness. We measured the septal thickness of ammonoid phragmocones through ontogeny in order to test the hypothesis that atmospheric pCO2, seawater pH and other factors affected aragonite biomineralisation in ammonoids. Particularly, we studied septal thickness of ammonoids before and after the ocean acidification event in the latest Triassic until the Early Cretaceous. Early Jurassic ammonoid lineages had thinner septa relative to diameter than their Late Triassic relatives, which we tentatively interpret as consequence of a positive selection for reduced shell material as an evolutionary response to this ocean acidification event. This response was preserved within several lineages among the Early Jurassic descendants of these ammonoids. By contrast, we did not find a significant correlation between septal thickness and long-term atmospheric pCO2 or seawater pH, but we discovered a correlation with palaeolatitude. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-022-00246-2.

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.

Historical significance and taxonomic status of Ischyrodon meriani (Pliosauridae) from the Middle Jurassic of Switzerland.

Ischyrodon meriani is an obscure pliosaurid taxon established upon an exceptionally large tooth crown of a probable Callovian (Middle Jurassic) age that originates from Wölflinswil, Canton of Aargau, Switzerland. Despite being known for almost two centuries, the specimen remains poorly researched. Historically, I. meriani has been associated, or even considered conspecific, with Pliosaurus macromerus and Liopleurodon ferox. However, neither of the two hypotheses have been tested through detailed comparisons or using modern quantitative methods. Here, we redescribe the type of Ischyrodon meriani, illustrate it, and compare to teeth of thalassophonean pliosaurids, with special focus on Jurassic representatives of the clade. Multivariate analyses show close similarities to L. ferox but comparable structures to those of I. meriani, including a distinctive pattern of the apicobasal ridges, are also observable in some mid-Cretaceous brachauchenines from the 'Polyptychodon' assemblage of East and South East England. While it is likely that I. meriani represents a Liopleurodon-like taxon, or is indeed conspecific with L. ferox, which would make I. meriani the proper name for the species, any such taxonomic considerations are hindered by the fragmentary nature of the type specimens of both these taxa as well as limited knowledge of the dental variability within and between individual jaws of L. ferox. Currently, I. meriani is best treated as a nomen dubium. Finally, we discuss the potential implications of I. meriani being conspecific with L. ferox, and additionally provide a commentary on the taxonomic status of Liopleurodon.

Early cephalopod evolution clarified through Bayesian phylogenetic inference.

BACKGROUND: Despite the excellent fossil record of cephalopods, their early evolution is poorly understood. Different, partly incompatible phylogenetic hypotheses have been proposed in the past, which reflected individual author's opinions on the importance of certain characters but were not based on thorough cladistic analyses. At the same time, methods of phylogenetic inference have undergone substantial improvements. For fossil datasets, which typically only include morphological data, Bayesian inference and in particular the introduction of the fossilized birth-death model have opened new possibilities. Nevertheless, many tree topologies recovered from these new methods reflect large uncertainties, which have led to discussions on how to best summarize the information contained in the posterior set of trees. RESULTS: We present a large, newly compiled morphological character matrix of Cambrian and Ordovician cephalopods to conduct a comprehensive phylogenetic analysis and resolve existing controversies. Our results recover three major monophyletic groups, which correspond to the previously recognized Endoceratoidea, Multiceratoidea, and Orthoceratoidea, though comprising slightly different taxa. In addition, many Cambrian and Early Ordovician representatives of the Ellesmerocerida and Plectronocerida were recovered near the root. The Ellesmerocerida is para- and polyphyletic, with some of its members recovered among the Multiceratoidea and early Endoceratoidea. These relationships are robust against modifications of the dataset. While our trees initially seem to reflect large uncertainties, these are mainly a consequence of the way clade support is measured. We show that clade posterior probabilities and tree similarity metrics often underestimate congruence between trees, especially if wildcard taxa are involved. CONCLUSIONS: Our results provide important insights into the earliest evolution of cephalopods and clarify evolutionary pathways. We provide a classification scheme that is based on a robust phylogenetic analysis. Moreover, we provide some general insights on the application of Bayesian phylogenetic inference on morphological datasets. We support earlier findings that quartet similarity metrics should be preferred over the Robinson-Foulds distance when higher-level phylogenetic relationships are of interest and propose that using a posteriori pruned maximum clade credibility trees help in assessing support for phylogenetic relationships among a set of relevant taxa, because they provide clade support values that better reflect the phylogenetic signal.

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.

The swimming trace Undichna from the latest Devonian Hangenberg Sandstone equivalent of Morocco.

Trace fossils occur in several strata of the Devonian and Carboniferous of the eastern Anti-Atlas, but they are still poorly documented. Here, we describe a fossil swimming trace from strata overlying the Hangenberg Black Shale (correlation largely based on lithostratigraphy; Postclymenia ammonoid genozone, ca. 370 Ma old). We discuss the systematic position of the tracemaker and its body size. This ichnofossil is important for three main reasons: (1) it is considered here to be the first record of Undichna from the Devonian of Gondwana, as far as we know; (2) it is the oldest record of vertebrate trace fossils from Africa; (3) it provides a unique window into the behaviour of Late Devonian fishes for which body-fossils cannot provide direct evidence. Further, we put this discovery into the macroecological context of the palaeoenvironment following the Late Devonian Hangenberg biodiversity crisis.

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.

A late-surviving stem-ctenophore from the Late Devonian of Miguasha (Canada).

Like other soft-bodied organisms, ctenophores (comb jellies) produce fossils only under exceptional taphonomic conditions. Here, we present the first record of a Late Devonian ctenophore from the Escuminac Formation from Miguasha in eastern Canada. Based on the 18-fold symmetry of this disc-shaped fossil, we assign it to the total-group Ctenophora. Our phylogenetic analyses suggest that the new taxon Daihuoides jakobvintheri gen. et sp. nov. falls near Cambrian stem ctenophores such as 'dinomischids' and 'scleroctenophorans'. Accordingly, Daihuoides is a Lazarus-taxon, which post-dates its older relatives by over 140 million years, and overlaps temporally with modern ctenophores, whose oldest representatives are known from the Early Devonian. Our analyses also indicate that the fossil record of ctenophores does not provide strong evidence for or against the phylogenomic hypothesis that ctenophores are sister to all other metazoans.