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 beargulchensisfrom 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.

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.

The fossilized birth–death (FBD) process provides an ideal model for inferring phylogenies from both extant and fossil taxa. Using this approach, fossils are directly integrated into the tree, leading to a statistically coherent prior on divergence times. Since fossils are typically not associated with molecular sequences, additional information is required to place fossils in the tree. We use simulations to evaluate two different approaches to handling fossil placement in FBD analyses: using topological constraints, where the user specifies monophyletic clades based on established taxonomy, or using total‐evidence analyses, which use a morphological data matrix in addition to the molecular alignment. We also explore how rate variation in fossil recovery or diversification rates impacts these approaches. We find that the extant topology is well recovered under all methods of fossil placement. Divergence times are similarly well recovered across all methods, with the exception of constraints which contain errors. We see similar patterns in datasets which include rate variation, however, relative errors in extant divergence times increase when more variation is included in the dataset, for all approaches using topological constraints, and particularly for constraints with errors. Finally, we show that trees recovered under the FBD model are more accurate than those estimated using non‐time calibrated inference. Overall, we show that both fossil placement approaches are reliable even when including uncertainty. Our results underscore the importance of core taxonomic research, including morphological data collection and species descriptions, irrespective of the approach to handling phylogenetic uncertainty using the FBD process.

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.

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.

Fluctuations in body size of orthoconic cephalopods are reported from late Silurian to Late Devonian sediments at several locations in the Tafilalt and in the Dra-Valley (Anti-Atlas, Morocco). The combination of measurements of diameters and apical angles allows the reconstruction of their total conch size (length and volume), which revealed a strongly right-skewed size distribution with an average length of 278 mm, while the largest Devonian actinocerids exceeded 2 m. Within the examined groups (Actinocerida, Orthocerida, Pseudorthocerida), there is no uniform trend, but rather frequent fluctuations with maximum sizes in the late Lochkovian and early Emsian. Body size decreased in times of extinction events, while stable periods are mostly associated with a size increase. Additionally, conch size correlates well with gamma diversity and global δ¹³C values. Furthermore, the apical angle and septal diameter of orthocones appear to correlate, but only when their mean values are compared across beds.