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The lack of any pterosaur living descendants creates gaps in the knowledge of the biology of this group, including its cervical biomechanics, which makes it difficult to understand their posture and life habits. To mitigate part of this issue, we reconstructed the cervical osteology and arthrology of three pterosaurs, allowing us to make inferences about the position of the neck of these animals at rest. We used scans of three-dimensionally preserved cervical series of Anhanguera piscator , Azhdarcho lancicollis and Rhamphorhynchus muensteri for the reconstructions, thus representing different lineages. For the recognition of ligaments, joint cartilages, and levels of overlapping of the zygapophyses, we applied the Extant Phylogenetic Bracket method, based on various extant birds and on Caiman latirostris . We inferred that pterosaur intervertebral joints were probably covered by a thin layer of synovial cartilage whose thickness varied along the neck, being thicker in the posterior region. Ignoring this cartilage can affect reconstructions. According to the vertebral angulation, their neck was slightly sinuous when in rest position. Our analyses also indicate that pterosaurs had segmented and supra-segmented articular cervical ligaments, which could confer stabilization, execute passive forces on the neck and store elastic energy.

The necks of pterosaurs were flexible and provided mobility for a relatively long skull. The varied morphologies and levels of pneumatization of their cervical vertebrae reflected differences in biomechanical behavior. Here, we examined the structural resistance of the cervical vertebrae to infer the most advantageous movements during the foraging behaviors of two pterodactyloid pterosaurs. We also examined the relationship between vertebral resistance and the presence of pneumatic foramina on the bone cortex. For this purpose, we analyzed three-dimensional models of the cervical vertebrae of Anhanguera piscator and Azhdarcho lancicollis , which are hypothesized to be aquatic and terrestrial predators, respectively, and employed Finite Element Analysis (FEA) to assess and quantify the stresses experienced by the vertebrae due to the performance of six different movement scenarios. We observed that the shorter vertebrae at the ends of the neck of both species favored the proliferation of larger stresses in these regions, especially in the posterior cervicals of Anhanguera piscator and in the atlas-axis of Azhdarcho lancicollis , and that their taller neural arches aided in absorbing stress. Larger stresses at the ends of the neck are consistent with the interior trabecular reinforcement of the atlas-axis and posterior cervical vertebrae, suggesting a link between biomechanical behavior and the level of pneumatization. Additionally, mechanical requirements may have also influenced the presence, size, and number of pneumatic foramina on the vertebral cortex, as evidenced by the large lateral foramen in Anhanguera piscator and the smaller and more numerous ones bordering the neural canal in Azhdarcho lancicollis . Our inferences corroborate the differences in foraging strategies hypothesized for anhanguerids and azhdarchids. The absorption of stresses resulting from ventral pitching of the head and neck indicates that the cervical vertebrae were well-adapted for making rapid movements during predatory hunting. However, variations in the height of the neural spine indicate different mechanical behaviors between these species when raising the skull and neck, which could be faster in Anhanguera piscator while more vigorous in Azhdarcho lancicollis .

Research on the postcranial skeletal pneumaticity in pterosaurs is common in the literature, but most studies present only qualitative assessments. When quantitative, they are done on isolated bones. Here, we estimate the Air Space Proportion (ASP) obtained from micro-CT scans of the sequence from the sixth cervical to the fourth dorsal vertebra of an anhanguerine pterosaur to understand how pneumaticity is distributed in these bones. Pneumatisation of the vertebrae varied between 68 and 72% of their total volume. The neural arch showed higher ASP in all vertebrae. Anhanguerine vertebral ASP was generally higher than in sauropod vertebrae but lower than in most extant birds. The ASP observed here is lower than that calculated for the appendicular skeleton of other anhanguerian pterosaurs, indicating the potential existence of variation between axial and appendicular pneumatisation. The results point to a pattern in the distribution of the air space, which shows an increase in the area occupied by the trabecular bone in the craniocaudal direction of the vertebral series and, in each vertebra, an increase of the thickness of the trabeculae in the zygapophyses. This indicates that the distribution of pneumatic diverticula in anhanguerine vertebrae may not be associated with stochastic patterns.

Mass mortality events are unusual in the Crato Formation. Although mayflies’ accumulations have been previously reported from that unit, they lacked crucial stratigraphic data. Here we provide the first taphonomic analysis of a mayfly mass mortality event, from a layer 285 cm from the top of the Formation, with 40 larvae, and an overview of the general biological community structure of a three meters deep excavated profile. The only other autochthonous taxon observed in the mayfly mortality layer was the gonorynchiform fish Dastilbe . The larvae and fishes were smaller than usual in the layer 285 cm, suggesting that they lived in a shallow water column. Their excellent preservation and a lack of preferential orientation in the samples suggest an absence of significant transport. All mayflies belong to the Hexagenitidae, whose larvae lived in quiet waters. We also recovered allochthonous taxa in that layer indicative of drier weather conditions. Adjacent layers presented crystals and pseudomorphs of halite, suggesting drought and high salinity. In other layers, Dastilbe juveniles were often found in mass mortality events, associated with a richer biota. Our findings support the hypothesis that the Crato Formation’s palaeolake probably experienced seasonal high evaporation, caused by the hot climate tending to aridity, affecting the few autochthonous fauna that managed to live in this setting.

Anhanguerids comprise an important clade of pterosaurs, mostly known from dozens of three-dimensionally preserved specimens recovered from the Lower Cretaceous Romualdo Formation (northeastern Brazil). They are remarkably diverse in this sedimentary unit, with eight named species, six of them belonging to the genus . However, such diversity is likely overestimated, as these species have been historically diagnosed based on subtle differences, mainly based on the shape and position of the cranial crest. In spite of that, recently discovered pterosaur taxa represented by large numbers of individuals, including juveniles and adults, as well as presumed males and females, have crests of sizes and shapes that are either ontogenetically variable or sexually dimorphic. We describe in detail the skull of one of the most complete specimens referred to , AMNH 22555, and use it as a case study to review the diversity of anhanguerids from the Romualdo Formation. In order to accomplish that, a geometric morphometric analysis was performed to assess size-dependent characters with respect to the premaxillary crest in the 12 most complete skulls bearing crests that are referred in, or related to, this clade, almost all of them analyzed first hand. Geometric morphometric regression of shape on centroid size was highly statistically significant (  = 0.0091) and showed that allometry accounts for 25.7% of total shape variation between skulls of different centroid sizes. Premaxillary crests are both taller and anteroposteriorly longer in larger skulls, a feature consistent with ontogenetic growth. A new diagnosis is proposed for , including traits that are nowadays known to be widespread within the genus, as well as ontogenetic changes. AMNH 22555 cannot be referred to \" \" and, in fact, \" \", \" \", and \" \" are here considered . Historically, minor differences in crest morphology have been used in the definition of new anhanguerid species. Nowadays, this practice resulted in a considerable difficulty in referring well-preserved skulls into known taxa. When several specimens are analyzed, morphologies previously believed to be disparate are, in fact, separated by a continuum, and are thus better explained as individual or temporal variations. Stratigraphically controlled excavations on the Romualdo Formation have showed evidence for faunal turnover regarding fish communities. It is thus possible that some of the pterosaurs from this unit were not coeval, and might even represent anagenetic morphotypes. Unfortunately, amateur collecting of Romualdo Formation fossils, aimed especially at commerce, resulted in the lack of stratigraphic data of virtually all its pterosaurs and precludes testing of these further hypotheses.

The Crato Formation (Aptian, Lower Cretaceous) is a fossiliferous deposit of global significance, representing a lacustrine palaeoenvironment which offers insights into aquatic insect taphonomy. Despite its importance, prior studies lacked an actualistic approach. Here, we analyze the preservation of mayflies (Ephemeroptera) and dragonflies (Odonata) from this formation using experimental taphonomy on 253 extant Ephemeroptera and 236 Odonata, alongside 306 fossil specimens. Disarticulation experiments showed that the thorax of modern mayfly larvae disarticulated first, yet Crato Hexagenitidae larvae retained intact thoraces, indicating minimal disturbance and autochthonous deposition. Fossil alate specimens rarely exhibited decay-related wing damage, aligning with short decay times. Dragonfly carcasses exhibited a characteristic leg posture in death, also preserved in Crato fossils, further suggesting minimal transport. Additionally, fossil dragonflies retained labial masks, the first structure to disarticulate experimentally, consistent with parautochthonous assemblages. Mayfly larvae exposed to low salinity during experiments exhibited excessive defecation before death, hinting at possible low salinity conditions in the Crato palaeoenvironment, though preservational challenges obscure confirmation. During experimentation, we also noticed that all carcasses immediately floated under hypersaline conditions, while carcasses immersed in non-hypersaline conditions went through slower decomposition. Thus, we can safely propose with experimental data that microbial biofilms on the surface of the water were acting during carcass sinking in this deposit.

Birds and pterosaurs have pneumatic bones, a feature likely related to their flight capabilities but whose evolution and origin is still poorly understood. Pneumatic foramina are present on the external surface of the bone and are reliable indicators of post-cranial skeletal pneumatization present in Pterosauria, Eusauropoda, and Neotheropoda. Here, we carried out a qualitative analysis of the position, size and number of pneumatic foramina of the cervical and thoracic/dorsal vertebrae of pterosaurs and birds, as they have the potential to challenge hypotheses about the emergence and evolution of the respiratory trait in these groups. We also discussed differences between pneumatic and vascular foramina for identification purposes. Besides phylogenetic representativeness, the pterosaur taxonomic sampling considered the preservation of specimens and, for birds, their life habit, as this relates to the level of pneumatization. Pneumatic foramina on the lateral faces of the centrum of the mid-cervical vertebrae of pterosaurs and birds differ in position and size, and those adjacent to the neural canal additionally differ in number. The avian posterior cervical vertebrae show a higher number of pneumatic foramina in comparison to their mid-cervicals, while the opposite is true for pterosaurs, suggesting differences in the cervical air sac of these clades. Pneumatic foramina were found at the base of the transverse processes of the notarial vertebrae of birds, while they were absent from some of the pterosaurs analyzed here, revealing the presence of a pneumatic hiatus in the vertebral column that might be explained due to the distance of this structure to the cervical air sac. These findings indicate that, although the overall skeletal pneumatization of pterosaurs and birds present deep homologies, some pneumatic features occurred convergently because variation in the number of pneumatic foramina along the vertebral column is related to the position of the air sacs in pterosaurs and birds and/or the habit of each species. There is an evident reduction of the pneumatic foramina in birds that have aquatic foraging and an increase in the ones which perform static soaring. Although we did not find any external anatomical difference between pneumatic and vascular foramina, we observed that vascular foramina occur at specific sites and thus identification on the basis of location is reliable.

Fossil eggs and embryos that provide unique information about the reproduction and early growth of vertebrates are exceedingly rare, particularly for pterosaurs. Here we report on hundreds of three-dimensional (3D) eggs of the species Hamipterus tianshanensis from a Lower Cretaceous site in China, 16 of which contain embryonic remains. Computed tomography scanning, osteohistology, and micropreparation reveal that some bones lack extensive ossification in potentially late-term embryos, suggesting that hatchlings might have been flightless and less precocious than previously assumed. The geological context, including at least four levels with embryos and eggs, indicates that this deposit was formed by a rare combination of events, with storms acting on a nesting ground. This discovery supports colonial nesting behavior and potential nesting site fidelity in the Pterosauria.

The pterosaur record from the Iberian Peninsula is mostly scarce and undefined, but in the last few years some new taxa have been described from different Lower Cretaceous sites of Spain. Here we describe a new genus and species of toothed pterodactyloid pterosaur from the Barremian of the Iberian Peninsula, Iberodactylus andreui gen. et sp. nov., that shows a close and rather unexpected relationship with Hamipterus tianshanensis from China. A review of the phylogenetic relationships of the Anhangueria reveals a new family of pterodactyloid pterosaurs, the Hamipteridae fam. nov. being recovered as sister-group of the Anhangueridae. This latter clade can be in turn divided into the new clades Anhanguerinae and Coloborhynchinae. The close relationships of Iberodactylus and Hamipterus shows an interesting palaeobiogeographical correlation between the Chinese and Iberian pterosaur faunas during the Barremian (Lower Cretaceous). The discovery of Iberodactylus strongly suggests that the clade Anhangueria has clear ancestral ties in eastern Laurasia.

Over a decade after the last major review of the Cambridge Greensand pterosaurs, their systematics remains one of the most disputed points in pterosaur taxonomy. Ornithocheiridae is still a wastebasket for fragmentary taxa, and some nomenclatural issues are still a problem. Here, the species from the Cretaceous of England that, at some point, were referred in Ornithocheirus, are reviewed. Investigation of the primary literature confirmed that Criorhynchus should be considered an objective junior synonym of Ornithocheirus. Taxonomic review of more than 30 species known from fragmentary remains showed that 16 of them are undiagnosable (nomina dubia): Palaeornis cliftii, Cimoliornis diomedeus, Pterodactylus compressirostris, Pterodactylus fittoni, Pterodactylus woodwardi, Ornithocheirus brachyrhinus, Ornithocheirus carteri, Ornithocheirus crassidens, Ornithocheirus dentatus, Ornithocheirus enchorhynchus, Ornithocheirus eurygnathus, Ornithocheirus oxyrhinus, Ornithocheirus scaphorhynchus, Ornithocheirus tenuirostris, Ornithocheirus xyphorhynchus, and Pterodactylus sagittirostris. Fourteen species are considered valid, and diagnoses are provided to all of them: Ornithocheirus simus, Lonchodraco giganteus comb. n., Lonchodraco machaerorhynchus comb. n., Lonchodraco(?) microdon comb. n., Coloborhynchus clavirostris, 'Ornithocheirus' capito, Camposipterus nasutus comb. n., Camposipterus(?) sedgwickii comb. n., Camposipterus(?) colorhinus comb. n., Cimoliopterus cuvieri comb. n., 'Ornithocheirus' polyodon, 'Ornithocheirus' platystomus, 'Pterodactylus' daviesii, and 'Ornithocheirus' denticulatus. These species are referred in the genera Ornithocheirus, Lonchodraco gen. n., Coloborhynchus, Cimoliopterus gen. n., and Camposipterus gen. n., but additional genera are probably present, as indicated by the use of single quotation marks throughout the text. A cladistic analysis demonstrates that Anhangueridae lies within a newly recognized clade, here named Anhangueria, which also includes the genera Cearadactylus, Brasileodactylus, Ludodactylus, and Camposipterus. The anhanguerian 'Cearadactylus' ligabuei belongs to a different genus than Cearadactylus atrox. Lonchodraconidaefam. n. (more or less equivalent to LonchodectidaesensuUnwin 2001) is a monophyletic entity, but its exact phylogenetic position remains uncertain, as is the case of Ornithocheirus simus. Therefore, it is proposed that Ornithocheiridae should be constricted to its type species and thus is redundant. Other taxa previously referred as \"ornithocheirids\" are discussed in light of the revised taxonomy.