The pseudosuchian record in paleohistology: A small review.

Archosauria originated around the Earth's largest biotic crisis that severely affected all ecosystems globally, the Permotriassic Mass extinction event, and comprises two crown-group lineages: the bird-lineage and the crocodylian lineage. The bird lineage includes the iconic pterosaurs, as well as dinosaurs and birds, whereas the crocodylian lineage includes clades such as aetosaurs, poposaurs, "rauisuchians," as well as Crocodylomorpha; the latter being represented today only by less than 30 extant species of Crocodylia. Despite playing important roles during Mesozoic and Cenozoic ecosystems, both on land and in water, Pseudosuchia received far less attention compared to the bird-lineage, which is also reflected in number and scope of histological studies so far. Lately, the field has seen a shift of focus toward pseudosuchians, however, and the symposium on "Paleohistological Inferences of Paleobiological Traits in Pseudosuchia" held during the International Congress of Vertebrate Morphology 2023 in Cairns, Queensland, Australia, is the latest proof of that. To put these novel aspects of paleohistological and paleobiological research into context, an overview of the non-extant pseudosuchian taxa whose postcranial bones were studied so far is provided here (c. 80 species out of a total of more than 700 extinct species described) and recent trends in pseudosuchian osteohistology are highlighted. In addition, histological studies on cranial and dental material and other potential hard tissues, such as eggshells and otoliths, are briefly reviewed as well.

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.

Asymmetric fin shape changes swimming dynamics of ancient marine reptiles' soft robophysical models.

Animals have evolved highly effective locomotion capabilities in terrestrial, aerial, and aquatic environments. Over life's history, mass extinctions have wiped out unique animal species with specialized adaptations, leaving paleontologists to reconstruct their locomotion through fossil analysis. Despite advancements, little is known about how extinct megafauna, such as the Ichthyosauria one of the most successful lineages of marine reptiles, utilized their varied morphologies for swimming. Traditional robotics struggle to mimic extinct locomotion effectively, but the emerging soft robotics field offers a promising alternative to overcome this challenge. This paper aims to bridge this gap by studyingMixosauruslocomotion with soft robotics, combining material modeling and biomechanics in physical experimental validation. Combining a soft body with soft pneumatic actuators, the soft robotic platform described in this study investigates the correlation between asymmetrical fins and buoyancy by recreating the pitch torque generated by extinct swimming animals. We performed a comparative analysis of thrust and torque generated byCarthorhyncus,Utatsusaurus,Mixosaurus,Guizhouichthyosaurus, andOphthalmosaurustail fins in a flow tank. Experimental results suggest that the pitch torque on the torso generated by hypocercal fin shapes such as found in model systems ofGuizhouichthyosaurus,MixosaurusandUtatsusaurusproduce distinct ventral body pitch effects able to mitigate the animal's non-neutral buoyancy. This body pitch control effect is particularly pronounced inGuizhouichthyosaurus, which results suggest would have been able to generate high ventral pitch torque on the torso to compensate for its positive buoyancy. By contrast, homocercal fin shapes may not have been conducive for such buoyancy compensation, leaving torso pitch control to pectoral fins, for example. Across the range of the actuation frequencies of the caudal fins tested, resulted in oscillatory modes arising, which in turn can affect the for-aft thrust generated.

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.

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.

Complex dental wear analysis reveals dietary shift in Triassic placodonts (Sauropsida, Sauropterygia).

Placodonts were durophagous reptiles of the Triassic seas with robust skulls, jaws, and enlarged, flat, pebble-like teeth. During their evolution, they underwent gradual craniodental changes from the Early Anisian to the Rhaetian, such as a reduction in the number of teeth, an increase in the size of the posterior palatal teeth, an elongation of the premaxilla/rostrum, and a widening of the temporal region. These changes are presumably related to changes in dietary habits, which, we hypothesise, are due to changes in the type and quality of food they consumed. In the present study, the dental wear pattern of a total of nine European Middle to Late Triassic placodont species were investigated using 2D and 3D microwear analyses to demonstrate whether there could have been a dietary shift or grouping among the different species and, whether the possible changes could be correlated with environmental changes affecting their habitats. The 3D analysis shows overlap between species with high variance between values and there is no distinct separation. The 2D analysis has distinguished two main groups. The first is characterised by low number of wear features and high percentage of large pits. The other group have a high feature number, but low percentage of small pits. The 2D analysis showed a correlation between the wear data and the size of the enlarged posterior crushing teeth. Teeth with larger sizes showed less wear feature (with higher pit ratio) but larger individual features. In contrast, the dental wear facet of smaller crushing teeth shows more but smaller wear features (with higher scratch number). This observation may be related to the size of the food consumed, i.e., the wider the crown, the larger food it could crush, producing larger features. Comparison with marine mammals suggests that the dietary preference of Placochelys, Psephoderma and Paraplacodus was not exclusively hard, thick-shelled food. They may have had a more mixed diet, similar to that of modern sea otters. The diet of Henodus may have included plant food, similar to the modern herbivore marine mammals and lizards. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-024-00304-x.

To glide or to swim? A reinvestigation of the enigmatic Wapitisaurus problematicus (Reptilia) from the Early Triassic of British Columbia, Canada.

Wapitisaurus problematicus was initially described as a member of the Weigeltisauridae, a clade of Late Permian gliding reptiles from Eurasia and Madagascar. However, the poor preservation of the holotype and only known specimen, from the lower Sulphur Mountain Formation at Ganoid Ridge (British Columbia, Canada), raised doubts about this assignment. Here, we redescribe W. problematicus and reassess its systematic position among diapsid reptiles. Comparison with all known weigeltisaurids, as well as contemporaneous reptiles from the Sulphur Mountain Formation, indicates that the taxon instead represents a thalattosauroid thalattosauriform, with noted similarities to Thalattosaurus and Paralonectes. This reidentification restricts weigeltisaurids to the Late Permian, with no occurrence in North America. Wapitisaurus problematicus potentially represents one of the oldest thalattosauriforms and increases our understanding of their diversity and disparity during the late Early and Middle Triassic. The close morphological similarities with later (thalattosauroid) thalattosauriforms and their high abundance in (shallow) marine settings may indicate an earlier invasion of this realm than previously assumed. This parallels observations in early ichthyopterygians with widespread opportunistic trophic niche diversification occurring relatively rapidly after the end-Permian mass extinction event.

Ontogenetic variation in the cranium of Mixosaurus cornalianus, with implications for the evolution of ichthyosaurian cranial development.

Relatively complete ontogenetic series are comparatively rare in the vertebrate fossil record. This can create biases in our understanding of morphology and evolution, since immaturity can represent a source of unrecognized intraspecific variation in both skeletal anatomy and ecology. In the extinct marine reptile clade Ichthyopterygia, ontogenetic series were widely studied only in some Jurassic genera, while the ontogeny of the oldest and most basal members of the clade is very poorly understood. Here, we investigate cranial ontogeny in Mixosaurus cornalianus, from the Middle Triassic Besano Formation of the Swiss and Italian Alps. This small-bodied taxon is represented by a wealth of material from multiple size classes, including fetal material. This allows us to assess ontogenetic changes in cranial morphology, and identify stages in the ontogenetic trajectory where divergence with more derived ichthyosaurs has occurred. Early ontogenetic stages of Mixosaurus show developmental patterns that are reminiscent of the presumed ancestral (early diverging sauropsid) condition. This is prominently visible in the late fetal stage in both the basioccipital, which shows morphology akin to basal tubera, and in the postorbital, which has a triradiate head. The ontogenetic trajectory of at least some of the cranial elements of Mixosaurus is therefore likely still very akin to the ancestral condition, even though the adult cranium diverges from the standard diapsid morphology. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00289-z.

High phenotypic plasticity at the dawn of the eosauropterygian radiation.

The initial radiation of Eosauropterygia during the Triassic biotic recovery represents a key event in the dominance of reptiles secondarily adapted to marine environments. Recent studies on Mesozoic marine reptile disparity highlighted that eosauropterygians had their greatest morphological diversity during the Middle Triassic, with the co-occurrence of Pachypleurosauroidea, Nothosauroidea and Pistosauroidea, mostly along the margins of the Tethys Ocean. However, these previous studies quantitatively analysed the disparity of Eosauropterygia as a whole without focussing on Triassic taxa, thus limiting our understanding of their diversification and morphospace occupation during the Middle Triassic. Our multivariate morphometric analyses highlight a clearly distinct colonization of the ecomorphospace by the three clades, with no evidence of whole-body convergent evolution with the exception of the peculiar pistosauroid Wangosaurus brevirostris, which appears phenotypically much more similar to nothosauroids. This global pattern is mostly driven by craniodental differences and inferred feeding specializations. We also reveal noticeable regional differences among nothosauroids and pachypleurosauroids of which the latter likely experienced a remarkable diversification in the eastern Tethys during the Pelsonian. Our results demonstrate that the high phenotypic plasticity characterizing the evolution of the pelagic plesiosaurians was already present in their Triassic ancestors, casting eosauropterygians as particularly adaptable animals.

Palaeobiological inferences of "rauisuchians" Fasolasuchus tenax (Los Colorados Fm., Argentina) and Prestosuchus chiniquensis (Santa Maria Super sequence, Brazil) from the Middle-Upper Triassic of South America based on microstructural analyses.

"Rauisuchia" is a non-monophyletic group of quadrupedal and carnivorous pseudosuchians that inhabited the entire world during the Middle-Upper Triassic period (Anisian/Ladinian-Rhaetian). In South America, "rauisuchians" reached the largest sizes among continental carnivores. Despite their important ecological role, some aspects of their palaeobiology have been poorly examined. Here, we study appendicular bones, dorsal ribs and osteoderms of two genera, the Argentinean Fasolasuchus tenax (PVL 3850, holotype) and the Brazilian Prestosuchus chiniquensis (SNSB-BSPG AS XXV) respectively. The femur of F. tenax is formed by laminar fibrolamellar bone, which is composed of non-fully monorefringent woven-fibred matrix and primary osteons; the dorsal rib has a Haversian bone composition with an external fundamental system recorded and the osteoderm is formed by well-organised parallel-fibred bone. The femur, humerus and fibula of P. chiniquensis are mostly composed of strongly arranged parallel-fibred bone and a laminar vascularisation. The minimal ages obtained correspond to 9 years for F. tenax (based on the maximum number of growth marks in the osteoderm) and 4 years for P. chiniquensis (obtained from the highest count of growth marks in the femur and in the humerus). F. tenax attained somatic and skeletal maturity, while P. chiniquensis was near to reaching skeletal and sexual maturity, but it was somatically immature. The overall rapid growth rate and the high and uniform vascularisation seems to imply that these features are common in most of "rauisuchians", except in P. chiniquensis.

A forged 'chimera' including the second specimen of the protostegid sea turtle Santanachelysgaffneyi and shell parts of the pleurodire Araripemys from the Lower Cretaceous Santana Group of Brazil.

Fossils of Cretaceous sea turtles adapted to an open marine lifestyle remain rare finds to date. Furthermore, the relationships between extant sea turtles, chelonioids, and other Mesozoic marine turtles are still contested, with one key species being Santanachelys gaffneyi Hirayama, 1998, long considered the earliest true sea turtle. The species is an Early Cretaceous member of Protostegidae, a controversial clade either placed within or closely related to Chelonioidea or, alternatively, along the stem lineage of hidden-neck turtles (Cryptodira) and representing an independent open marine radiation. Santanachelys gaffneyi is one of the most completely preserved early protostegids and is therefore critical for establishing the global phylogenetic position of the group. However, the single known specimen of this taxon is yet to be described in detail. Here we describe a second specimen of Santanachelys gaffneyi from its type horizon, the Romualdo Formation (late Aptian) of the Santana Group of the Araripe basin, NE Brazil. The skeletal elements preserved include the posterior part of the skull, neck vertebrae, shoulder girdle, anterior-most and left/central part of the carapace with few peripherals, and plastron lacking most of the hyoplastra. The remaining part of the carapace was apparently completed by fossil dealers using an anterior part of the pleurodiran Araripemydidae, tentatively identified as a shell portion of cf. Araripemys barretoi, a more common Santana fossil turtle, among other indeterminate turtle shell fragments. The purpose of this paper is to report the repatriation of the specimen to Brazil and to provide a preliminary description. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00271-9.

Semicircular canal shape diversity among modern lepidosaurs: life habit, size, allometry.

BACKGROUND: The shape of the semicircular canals of the inner ear of living squamate reptiles has been used to infer phylogenetic relationships, body size, and life habits. Often these inferences are made without controlling for the effects of the other ones. Here we examine the semicircular canals of 94 species of extant limbed lepidosaurs using three-dimensional landmark-based geometric morphometrics, and analyze them in phylogenetic context to evaluate the relative contributions of life habit, size, and phylogeny on canal shape. RESULTS: Life habit is not a strong predictor of semicircular canal shape across this broad sample. Instead, phylogeny plays a major role in predicting shape, with strong phylogenetic signal in shape as well as size. Allometry has a limited role in canal shape, but inner ear size and body mass are strongly correlated. CONCLUSIONS: Our wide sampling across limbed squamates suggests that semicircular canal shape and size are predominantly a factor of phylogenetic relatedness. Given the small proportion of variance in semicircular canal shape explained by life habit, it is unlikely that unknown life habit could be deduced from semicircular canal shape alone. Overall, semicircular canal size is a good estimator of body length and even better for body mass in limbed squamates. Semiaquatic taxa tend to be larger and heavier than non-aquatic taxa, but once body size and phylogeny are accounted for, they are hard to distinguish from their non-aquatic relatives based on bony labyrinth shape and morphology.

A new long-necked archosauromorph from the Guanling Formation (Anisian, Middle Triassic) of southwestern China and its implications for neck evolution in tanystropheids.

A long neck is an evolutionary innovation convergently appearing in multiple tetrapod lineages, including groups of plesiosaurs, non-archosauriform archosauromorphs, turtles, sauropodomorphs, birds, and mammals. Among all tetrapods both extant and extinct, two Triassic archosauromorphs, Tanystropheus and Dinocephalosaurus, have necks that are particularly elongated relative to the lengths of their trunks. However, the evolutionary history of such hyper-elongated necks in these two archosauromorph clades remains unknown, partially because known close relatives such as Macrocnemus and Pectodens possess only moderately elongated necks. Here, we describe a newly discovered early diverging archosauromorph, Gracilicollum latens gen. et sp. nov., based on a specimen comprising a partial neck and an incompletely preserved skull. The long neck is composed of at least 18 cervical vertebrae. The dentition suggests that this new taxon most likely represents an aquatic piscivore, similar to Dinocephalosaurus and Tanystropheus hydroides. Despite possessing a high number of cervical vertebrae, Gracilicollum gen. nov. is recovered as a tanystropheid in an evolutionary grade between Macrocnemus and Tanystropheus rather than as a close relative of Dinocephalosaurus, a result that is primarily attributable to the presence of palatal teeth and the anatomy of the cervical vertebrae in Gracilicollum gen. nov. Considering the information provided by the new specimen, we provide a detailed discussion of the cervical evolution in dinocephalosaurids and tanystropheids, which is shown to be highly complex and mosaic in nature.

Heads or tails first? Evolution of fetal orientation in ichthyosaurs, with a scrutiny of the prevailing hypothesis.

According to a longstanding paradigm, aquatic amniotes, including the Mesozoic marine reptile group Ichthyopterygia, give birth tail-first because head-first birth leads to increased asphyxiation risk of the fetus in the aquatic environment. Here, we draw upon published and original evidence to test two hypotheses: (1) Ichthyosaurs inherited viviparity from a terrestrial ancestor. (2) Asphyxiation risk is the main reason aquatic amniotes give birth tail-first. From the fossil evidence, we conclude that head-first birth is more prevalent in Ichthyopterygia than previously recognized and that a preference for tail-first birth likely arose in derived forms. This weakens the support for the terrestrial ancestry of viviparity in Ichthyopterygia. Our survey of extant viviparous amniotes indicates that fetal orientation at birth reflects a broad diversity of factors unrelated to aquatic vs. terrestrial habitat, further undermining the asphyxiation hypothesis. We propose that birth preference is based on parturitional mechanics or carrying efficiency rather than habitat.

Unique bone microanatomy reveals ancestry of subterranean specializations in mammals.

Acquiring a subterranean lifestyle entails a substantial shift for many aspects of terrestrial vertebrates' biology. Although this lifestyle is associated with multiple instances of convergent evolution, the relative success of some subterranean lineages largely remains unexplained. Here, we focus on the mammalian transitions to life underground, quantifying bone microanatomy through high-resolution X-ray tomography. The true moles stand out in this dataset. Examination of this family's bone histology reveals that the highly fossorial moles acquired a unique phenotype involving large amounts of compacted coarse cancellous bone. This phenotype exceeds the adaptive optimum seemingly shared by several other subterranean mammals and can be traced back to some of the first known members of the family. This remarkable microanatomy was acquired early in the history of the group and evolved faster than the gross morphology innovations of true moles' forelimb. This echoes the pattern described for other lifestyle transitions, such as the acquisition of bone mass specializations in secondarily aquatic tetrapods. Highly plastic traits-such as those pertaining to bone structure-are hence involved in the early stages of different types of lifestyle transitions.

First evidence of Proganochelys quenstedtii (Testudinata) from the Plateosaurus bonebeds (Norian, Late Triassic) of Frick, Canton Aargau, Switzerland.

Proganochelys quenstedtii represents the best-known stem turtle from the Late Triassic, with gross anatomical and internal descriptions of the shell, postcranial bones and skull based on several well-preserved specimens from Central European fossil locations. We here report on the first specimen of P. quenstedtii from the Late Triassic (Klettgau Formation) Frickberg near the town of Frick, Canton Aargau, Switzerland. Similar to other Late Triassic 'Plateosaurus-bearing bonebeds', Proganochelys is considered to be a rare faunal element in the Swiss locality of Frick as well. The specimen, which is largely complete but was found only partially articulated and mixed with large Plateosaurus bones, overall resembles the morphology of the classical specimens from Germany. Despite being disarticulated, most skull bones could be identified and micro-computed tomography (CT) scanning of the posterior skull region reveals new insights into the braincase and neurovascular anatomy, as well as the inner ear region. These include the presence of a fenestra perilymphatica, potentially elongated cochlear ducts, and intense vascularization of small tubercles on the posterior end of the skull roof, which we interpret as horn cores. Other aspects of the skull in the braincase region, such as the presence or absence of a supratemporal remain ambiguous due to the fusion of individual bones and thus lack of visible sutures (externally and internally). Based on the size of the shell and fusion of individual elements, the specimen is interpreted as a skeletally mature animal. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-022-00260-4.

A toolbox for the retrodeformation and muscle reconstruction of fossil specimens in Blender.

Accurate muscle reconstructions can offer new information on the anatomy of fossil organisms and are also important for biomechanical analysis (multibody dynamics and finite-element analysis (FEA)). For the sake of simplicity, muscles are often modelled as point-to-point strands or frustra (cut-off cones) in biomechanical models. However, there are cases in which it is useful to model the muscle morphology in three dimensions, to better examine the effects of muscle shape and size. This is especially important for fossil analyses, where muscle force is estimated from the reconstructed muscle morphology (rather than based on data collected in vivo). The two main aims of this paper are as follows. First, we created a new interactive tool in the free open access software Blender to enable interactive three-dimensional modelling of muscles. This approach can be applied to both palaeontological and human biomechanics research to generate muscle force magnitudes and lines of action for FEA. Second, we provide a guide on how to use existing Blender tools to reconstruct distorted or incomplete specimens. This guide is aimed at palaeontologists but can also be used by anatomists working with damaged specimens or to test functional implication of hypothetical morphologies.

A new pachypleurosaur from the Early Ladinian Prosanto Formation in the Eastern Alps of Switzerland.

The Alpine Prosanto Formation (Middle Triassic) cropping out in the Ducan region in eastern Switzerland has yielded a rich fish and reptile fauna. Here, we present new pachypleurosaur remains from the upper part of the formation (Early Ladinian), similar to the previously known pachypleurosaurs from the Middle Triassic UNESCO World Heritage Site of Monte San Giorgio in southern Switzerland/northern Italy. From these remains, a new pachypleurosaur species, Prosantosaurus scheffoldi nov. gen. et spec., is described on the basis of six fairly complete skeletons, one disarticulated specimen and an isolated skull. As is typical for pachypleurosaurs and most other Triassic marine reptiles, the new taxon is based to a large degree on a combination of characters (e.g., nasals articulating broadly with the anterior margins of the prefrontals and lacking posterior processes; postorbitals with rounded anterior processes that articulate with the postfrontals anterolaterally) rather than on many unambiguous autapomorphies, although a few of the latter were found including (1) a premaxilla which is excluded from entering both the external and internal nares and (2) a parietal, which is distinctly longer than wide and carrying distinct anterolaterally angled processes. Phylogenetic relationships of the new taxon are tested within European Pachypleurosauria, revealing that the new species is the sister taxon to a clade including Serpianosaurus, Proneusticosaurus, and the monophyletic Neusticosaurus spp. Mapping of palaeogeographic and stratigraphical distribution of valid European pachypleurosaurs shows that a formerly proposed scenario of migration of pachypleurosaurs from the eastern Palaeotethys during the Olenekian into the Germanic Basin and a subsequent diversification and invasion during the Anisian into the intraplatform basins of the South Alpine realm must be re-assessed. The exceptional preservation and preparation of the Ducan fossils further allow the description of tooth replacement patterns for the first time in a European pachypleurosaur species. The "alveolarization" of replacement teeth, the horizontal replacement pattern, and the subsequent remodelling of the functional alveoli during tooth replacement supports the monophyly of Sauropterygia as discussed before. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-022-00254-2.

Placodus (Placodontia, Sauropterygia) dentaries from Winterswijk, The Netherlands (middle Anisian) and Hünfeld, Hesse, Germany (late Anisian) with comments on ontogenetic changes.

Two recently found dentaries from the Lower Muschelkalk of Winterswijk (The Netherlands) and from the Upper Muschelkalk of an outcrop in the vicinity of Hünfeld (Hesse, Germany) are studied and compared to lower jaws of placodonts. As a result, the here described specimens can be assigned to Placodus cf. gigas. However, this assignment should be regarded as preliminary due to the isolated nature of the material. More diagnostic material is necessary to validate this affiliation. A certain morphological variability in P. gigas dentaries that had been pointed out before is also obvious among the new material. Placodus gigas has a wide paleogeography and stratigraphic range and a revision of the material assigned to P. gigas with new methods is overdue but beyond the scope of the current paper. The dentary from Hünfeld is with about 4 cm preserved length the smallest so far known dentary of a Placodus. It provides interesting insights in morphological changes during ontogeny and reveals differences in trajectories when compared to dentaries of different ontogenetic stages of Cyamodus hildegardis.

A large osteoderm-bearing rib from the Upper Triassic Kössen Formation (Norian/Rhaetian) of eastern Switzerland.

An important component of the Alpine vertebrate record of Late Triassic age derives from the Kössen Formation, which crops out extensively in the eastern Alps. Here, we present an isolated and only partially preserved large rib, which carries an osteoderm on a low uncinate process. Osteological comparison indicates that the specimen likely belongs to a small clade of marine reptiles, Saurosphargidae. Members of the clade are restricted to the western (today Europe) and eastern margins of the Tethys (today China) and were so far known only from the Anisian stage of the Middle Triassic. The assignment of the new find to cf. Saurosphargidae, with potential affinities to the genus Largocephalosaurus from the Guanling Formation of Yunnan and Guizhou Provinces, China, would extend the occurrence of the clade about 35 million years into the Late Triassic.