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.

In some Devonian strata in the eastern Anti‐Atlas, fossil invertebrates are abundant, display a high taxonomic diversity and indicate many shifts in palaeoecology. This is reflected in changes in faunal composition of invertebrates and vertebrates. Fossils of jawed vertebrates of late Lochkovian and younger age have been recorded and are relatively common with their abundance and diversity increasing towards the Late Devonian. Environmental changes in the Devonian also left their mark in the preservation of vertebrates and invertebrates from the Anti‐Atlas, which varies strongly through time and regionally. This variation partially reflects environmental changes linked with the evolution of small marine basins during the disintegration of the continental shelf of Gondwana in this region, fluctuations of the regional sea level and other environmental changes. To improve our understanding of these ecological changes, of shifts in preservation through the succession and of the formation of Fossil‐Lagerstätten, we analysed the mineral composition of some invertebrate and vertebrate samples of Devonian and Early Carboniferous age by Raman spectroscopy and X‐ray diffraction. Additionally, we characterized some of these Fossil‐Lagerstätten using palaeontological and sedimentological parameters. We examined eight Devonian Konzentrat‐Lagerstätten and two Konservat‐Lagerstätten with soft‐tissue preservation (the Famennian Thylacocephalan Layer and the Hangenberg Black Shale of the southern Maïder). The last two are the first Konservat‐Lagerstätten described from the Devonian of North Africa. The taphonomic and oceanic settings suggest that these Konservat‐Lagerstätten are formed because of stagnation (related to vertical restriction of water exchange and water depth rather than limited spatial water exchange and a lateral restriction) in the relatively small Maïder Basin with limited water exchange with the neighbouring Tafilalt Basin. The temporally low oxygen levels in the Maïder Basin are a possible reason for the reduced chondrichthyan diversity (missing demersal and shallow water species) compared to the Tafilalt Platform.

During the Palaeozoic, a diversification in modes of life occurred that included a wide range of predators. Major macroecological events include the Cambrian Explosion (including the Agronomic Substrate Revolution and the here introduced 'Ediacaran-Cambrian Mouthpart Armament'), the Great Ordovician Biodiversification Event, the Palaeozoic Plankton Revolution, the Siluro-Devonian Jaw Armament (newly introduced herein) and the Devonian Nekton Revolution. Here, we discuss the evolutionary advancement in oral equipment, i.e. the Palaeozoic evolution of mouthparts and jaws in a macroecological context. It appears that particularly the latest Neoproterozoic to Cambrian and the Silurian to Devonian were phases when important innovations in the evolution of oral structures occurred.•

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.