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The Velaux-La Bastide Neuve fossil-bearing site (Bouches-du-Rhône, France) has yielded a diverse vertebrate assemblage dominated by dinosaurs, including the titanosaur Atsinganosaurus velauciensis. We here provide a complete inventory of vertebrate fossils collected during two large-scale field campaigns. Numerous crocodilian teeth occur together with complete skulls. Pterosaur, hybodont shark and fish elements are also represented but uncommon. Magnetostratigraphic analyses associated with biostratigraphic data from dinosaur eggshell and charophytes suggest a Late Campanian age for the locality. Lithologic and taphonomic studies, associated with microfacies and palynofacies analyses, indicate a fluvial setting of moderate energy with broad floodplain. Palynomorphs are quite rare; only three taxa of pollen grains occur: a bisaccate taxon, a second form probably belonging to the Normapolles complex, and another tricolporate taxon. Despite the good state of preservation, these taxa are generally difficult to identify, since they are scarce and have a very minute size. Most of the vertebrate remains are well preserved and suggest transport of the carcasses over short distances before accumulation in channel and overbank facies, together with reworked Aptian grains of glauconite, followed by a rapid burial. The bones accumulated in three thin layers that differ by their depositional modes and their taphonomic histories. Numerous calcareous and iron oxides-rich paleosols developed on the floodplain, suggesting an alternating dry and humid climate in the region during the Late Campanian.

Feathers are amongst the most complex epidermal structures known and they have a well-documented evolutionary trajectory across non-avian dinosaurs and basal birds. Moreover, melanosome-like microbodies preserved in association with fossil plumage have been used to reconstruct original colour, behaviour and physiology. However, these putative ancient melanosomes might alternatively represent microorganismal residues, a conflicting interpretation compounded by a lack of unambiguous chemical data. We therefore used sensitive molecular imaging, supported by multiple independent analytical tests, to demonstrate that the filamentous epidermal appendages in a new specimen of the Jurassic paravian Anchiornis comprise remnant eumelanosomes and fibril-like microstructures, preserved as endogenous eumelanin and authigenic calcium phosphate. These results provide novel insights into the early evolution of feathers at the sub-cellular level and unequivocally determine that melanosomes can be preserved in fossil feathers.

Diverse epidermal appendages including grouped filaments closely resembling primitive feathers in non-avian theropods, are associated with skeletal elements in the primitive ornithischian dinosaur Kulindadromeus zabaikalicus from the Kulinda locality in south-eastern Siberia. This discovery suggests that “feather-like” structures did not evolve exclusively in theropod dinosaurs, but were instead potentially widespread in the whole dinosaur clade. The dating of the Kulinda locality is therefore particularly important for reconstructing the evolution of “feather-like” structures in dinosaurs within a chronostratigraphic framework. Here we present the first dating of the Kulinda locality, combining U-Pb analyses (LA-ICP-MS) on detrital zircons and monazites from sedimentary rocks of volcaniclastic origin and palynological observations. Concordia ages constrain the maximum age of the volcaniclastic deposits at 172.8 ± 1.6 Ma, corresponding to the Aalenian (Middle Jurassic). The palynological assemblage includes taxa that are correlated to Bathonian palynozones from western Siberia, and therefore constrains the minimum age of the deposits. The new U-Pb ages, together with the palynological data, provide evidence of a Bathonian age—between 168.3 ± 1.3 Ma and 166.1 ± 1.2 Ma—for Kulindadromeus . This is older than the previous Late Jurassic to Early Cretaceous ages tentatively based on local stratigraphic correlations. A Bathonian age is highly consistent with the phylogenetic position of Kulindadromeus at the base of the neornithischian clade and suggests that cerapodan dinosaurs originated in Asia during the Middle Jurassic, from a common ancestor that closely looked like Kulindadromeus . Our results consequently show that Kulindadromeus is the oldest known dinosaur with “feather-like” structures discovered so far.

Remarkably well-preserved soft tissues in Mesozoic fossils have yielded substantial insights into the evolution of feathers 1 . New evidence of branched feathers in pterosaurs suggests that feathers originated in the avemetatarsalian ancestor of pterosaurs and dinosaurs in the Early Triassic 2 , but the homology of these pterosaur structures with feathers is controversial 3 , 4 . Reports of pterosaur feathers with homogeneous ovoid melanosome geometries 2 , 5 suggest that they exhibited limited variation in colour, supporting hypotheses that early feathers functioned primarily in thermoregulation 6 . Here we report the presence of diverse melanosome geometries in the skin and simple and branched feathers of a tapejarid pterosaur from the Early Cretaceous found in Brazil. The melanosomes form distinct populations in different feather types and the skin, a feature previously known only in theropod dinosaurs, including birds. These tissue-specific melanosome geometries in pterosaurs indicate that manipulation of feather colour—and thus functions of feathers in visual communication—has deep evolutionary origins. These features show that genetic regulation of melanosome chemistry and shape 7 – 9 was active early in feather evolution. Melanosomes preserved in the skin and feathers of a tapejarid pterosaur from the Early Cretaceous found in Brazil provide evidence of the early use of feathers for visual communication.

A series of pterosaur bones from the Upper Cretaceous (Campanian) of Velaux (Bouches-du-Rhône, southeastern France) is described. This material, including both cranial and postcranial elements found in close association and likely belonging to a single immature individual, is assigned to a new genus and species of azhdarchid pterosaur, Mistralazhdarcho maggii. This large-sized taxon (wingspan ca. 4.5 m in the holotype, possibly reaching 5–6 m in mature individuals) is characterized by a slightly downturned mandibular symphysis that shows a ‘V’-shaped cross-sectional profile and bears a well-developed, anteriorly located median eminence on its dorsal surface. The presence of a median eminence suggests that Mistralazhdarcho might be closely related to Alanqa from the Cenomanian of Morocco. The material described here represents the first partial skeleton of a pterosaur recovered from the Late Cretaceous deposits of western Europe, and the new taxon is one of the most completely known European azhdarchids. Mistralazhdarcho is intermediate in size between the medium-sized genus Eurazhdarcho and the giant-sized genus Hatzegopteryx, two azhdarchids from the Maastrichtian of Romania. The discovery of Mistralazhdarcho suggests the presence of a third azhdarchid size class in the continental ecosystems of the latest Cretaceous European archipelago.

Rhabdodontidae is a successful clade of ornithopod dinosaurs, characteristic of Late Cretaceous continental faunas in Europe. A new rhabdodontid from the late Campanian, of southern France, Matheronodon provincialis gen. et sp. nov., is characterized by the extreme enlargement of both its maxillary and dentary teeth, correlated to a drastic reduction in the number of maxillary teeth (4 per generation in MMS/VBN-02-102). The interalveolar septa on the maxilla are alternately present or resorbed ventrally so as to be able to lodge such enlarged teeth. The rhabdodontid dentition and masticatory apparatus were adapted for producing a strict and powerful shearing action, resembling a pair of scissors. With their relatively simple dentition, contrasting with the sophisticated dental batteries in contemporary hadrosaurids, Matheronodon and other rhabdodontids are tentatively interpreted as specialized consumers of tough plant parts rich in sclerenchyma fibers, such as Sabalites and Pandanites .

With approximately 1,500 extant species, freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans. Nevertheless, their fossil record is extremely limited: only Potamidae, Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far. This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France), in close association with large vertebrate remains. In addition to (1) the systematic assignment of these claws, the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils, and (3) the likely scenario for their auto/allochthony in the Velaux fluvial system. These claws belong to a new taxon, Dinocarcinus velauciensis n. gen. n. sp., referred to as Portunoidea sensu lato, a group of “true” crabs nowadays linked to marine systems. However, the faunal assemblage, the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology, making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma. Either as primary or as secondary freshwater crabs, the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time, as early as the Campanian.

The posterior part of a skull from a tapejarid pterosaur was studied for the exceptional preservation of its soft-tissues. The specimen comes from the Early Cretaceous (Aptian) Crato Formation of northeastern Brazil. It shows a peculiar soft sagittal crest that gives new information about the anatomy and the diversity of tapejarids. The specimen lies on five limestone slabs. The bony part comprised the posterior part of the skull, with the braincase and one complex of three dorsally oriented and associated bony crests. The crest consists of a huge and elongated parietal crest, a trabecular frontal crest, and an occipital spine. The specimen preserves the posterior parts of the nasoantorbital fenestra and orbit. The soft cranial crest is characterized by a multilinear dorsal crest with almost parallel fibres, which curve posteriorly. Pigment organelles – melanosomes – were observed in the crest as well as ‘hair-like’ fibres (pycnofibres). They present diverse shapes and sizes, suggesting diverse colorations for the soft tissues. The new tapejarid shares certain features with the tapejarids Tupandactylus imperator and “Huaxiapterus” benxiensis, which include an elongated parieto-occipital crest. Nonetheless, it differs from Tupandactylus imperator and Ingridia navigans by the following characteristics: absence of a suprapremaxillary spine, very dorsally oriented posterior crests, as well as a specific morphology for the soft crest. Our specimen can therefore be assigned to a new taxon of Tapejaridae, a group previously reported from the Crato Formation.

Abstract With approximately 1,500 extant species, freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans. Nevertheless, their fossil record is extremely limited: only Potamidae, Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far. This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France), in close association with large vertebrate remains. In addition to (1) the systematic assignment of these claws, the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils, and (3) the likely scenario for their auto/allochtony in the Velaux fluvial system. These claws belong to a new taxon, Dinocarcinus velauciensis n. gen. n. sp., referred to as Portunoidea sensu lato, a group of “true” crabs nowadays linked to marine systems. However, the faunal assemblage, the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology, making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma. Either as primary or as secondary freshwater crabs, the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time, as early as the Campanian.