Pterosaurs were the first vertebrates to evolve powered flight and the largest animals to ever take wing. The pterosaurs persisted for over 150 million years before disappearing at the end of the Cretaceous, but the patterns of and processes driving their extinction remain unclear. Only a single family, Azhdarchidae, is definitively known from the late Maastrichtian, suggesting a gradual decline in diversity in the Late Cretaceous, with the Cretaceous-Paleogene (K-Pg) extinction eliminating a few late-surviving species. However, this apparent pattern may simply reflect poor sampling of fossils. Here, we describe a diverse pterosaur assemblage from the late Maastrichtian of Morocco that includes not only Azhdarchidae but the youngest known Pteranodontidae and Nyctosauridae. With 3 families and at least 7 species present, the assemblage represents the most diverse known Late Cretaceous pterosaur assemblage and dramatically increases the diversity of Maastrichtian pterosaurs. At least 3 families-Pteranodontidae, Nyctosauridae, and Azhdarchidae-persisted into the late Maastrichtian. Late Maastrichtian pterosaurs show increased niche occupation relative to earlier, Santonian-Campanian faunas and successfully outcompeted birds at large sizes. These patterns suggest an abrupt mass extinction of pterosaurs at the K-Pg boundary.
BACKGROUND: Pterosaurs have been known from the Cretaceous sediments of the Isle of Wight (southern England, United Kingdom) since 1870. We describe the three-dimensional pelvic girdle and associated vertebrae of a small near-adult pterodactyloid from the Atherfield Clay Formation (lower Aptian, Lower Cretaceous). Despite acknowledged variation in the pterosaur pelvis, previous studies have not adequately sampled or incorporated pelvic characters into phylogenetic analyses. METHODOLOGYPRINCIPAL FINDINGS: The new specimen represents the new taxon Vectidraco daisymorrisae gen. et sp. nov., diagnosed by the presence of a concavity posterodorsal to the acetabulum and the form of its postacetabular process on the ilium. Several characters suggest that Vectidraco belongs to Azhdarchoidea. We constructed a pelvis-only phylogenetic analysis to test whether the pterosaur pelvis carries a useful phylogenetic signal. Resolution in recovered trees was poor, but they approximately matched trees recovered from analyses of total evidence. We also added Vectidraco and our pelvic characters to an existing total-evidence matrix for pterosaurs. Both analyses recovered Vectidraco within Azhdarchoidea. CONCLUSIONSSIGNIFICANCE: The Lower Cretaceous strata of western Europe have yielded members of several pterosaur lineages, but Aptian pterosaurs from western Europe are rare. With a pelvis length of 40 mm, the new animal would have had a total length of c. 350 mm, and a wingspan of c. 750 mm. Barremian and Aptian pterodactyloids from western Europe show that small-bodied azhdarchoids lived alongside ornithocheirids and istiodactylids. This assemblage is similar in terms of which lineages are represented to the coeval beds of Liaoning, China; however, the number of species and specimens present at Liaoning is much higher. While the general phylogenetic composition of western European and Chinese communities appear to have been approximately similar, the differences may be due to different palaeoenvironmental and depositional settings. The western Europe pterodactyloid record may therefore be artificially low in diversity due to preservational factors.
Azhdarchid pterosaurs include the largest animals to ever take to the skies with some species exceeding 10 metres in wingspan and 220 kg in mass. Associated skeletons show that azhdarchids were long-necked, long-jawed predators that combined a wing planform suited for soaring with limb adaptations indicative of quadrupedal terrestrial foraging. The postcranial proportions of the group have been regarded as uniform overall, irrespective of their overall size, notwithstanding suggestions that minor variation may have been present. Here, we discuss a recently discovered giant azhdarchid neck vertebra referable to Hatzegopteryx from the Maastrichtian Sebeş Formation of the Transylvanian Basin, Romania, which shows how some azhdarchids departed markedly from conventional views on their proportions. This vertebra, which we consider a cervical VII, is 240 mm long as preserved and almost as wide. Among azhdarchid cervicals, it is remarkable for the thickness of its cortex (4-6 mm along its ventral wall) and robust proportions. By comparing its dimensions to other giant azhdarchid cervicals and to the more completely known necks of smaller taxa, we argue that Hatzegopteryx had a proportionally short, stocky neck highly resistant to torsion and compression. This specimen is one of several hinting at greater disparity within Azhdarchidae than previously considered, but is the first to demonstrate such proportional differences within giant taxa. On the assumption that other aspects of Hatzegopteryx functional anatomy were similar to those of other azhdarchids, and with reference to the absence of large terrestrial predators in the Maastrichtian of Transylvania, we suggest that this pterosaur played a dominant predatory role among the unusual palaeofauna of ancient Haţeg.
A pterosaur bone bed with at least 47 individuals (wing spans: 0.65-2.35 m) of a new species is reported from southern Brazil from an interdunal lake deposit of a Cretaceous desert, shedding new light on several biological aspects of those flying reptiles. The material represents a new pterosaur, Caiuajara dobruskii gen. et sp. nov., that is the southermost occurrence of the edentulous clade Tapejaridae (Tapejarinae, Pterodactyloidea) recovered so far. Caiuajara dobruskii differs from all other members of this clade in several cranial features, including the presence of a ventral sagittal bony expansion projected inside the nasoantorbital fenestra, which is formed by the premaxillae; and features of the lower jaw, like a marked rounded depression in the occlusal concavity of the dentary. Ontogenetic variation of Caiuajara dobruskii is mainly reflected in the size and inclination of the premaxillary crest, changing from small and inclined (∼115°) in juveniles to large and steep (∼90°) in adults. No particular ontogenetic features are observed in postcranial elements. The available information suggests that this species was gregarious, living in colonies, and most likely precocial, being able to fly at a very young age, which might have been a general trend for at least derived pterosaurs.
The pterosaurs were a diverse group of Mesozoic flying reptiles that underwent a body plan reorganization, adaptive radiation, and replacement of earlier forms midway through their long history, resulting in the origin of the Pterodactyloidea, a highly specialized clade containing the largest flying organisms. The sudden appearance and large suite of morphological features of this group were suggested to be the result of it originating in terrestrial environments, where the pterosaur fossil record has traditionally been poor [1, 2], and its many features suggested to be adaptations to those environments [1, 2]. However, little evidence has been available to test this hypothesis, and it has not been supported by previous phylogenies or early pterodactyloid discoveries. We report here the earliest pterosaur with the diagnostic elongate metacarpus of the Pterodactyloidea, Kryptodrakon progenitor, gen. et sp. nov., from the terrestrial Middle-Upper Jurassic boundary of Northwest China. Phylogenetic analysis confirms this species as the basalmost pterodactyloid and reconstructs a terrestrial origin and a predominantly terrestrial history for the Pterodactyloidea. Phylogenetic comparative methods support this reconstruction by means of a significant correlation between wing shape and environment also found in modern flying vertebrates, indicating that pterosaurs lived in or were at least adapted to the environments in which they were preserved.
A new species of pterosaur, Maaradactylus kellneri gen. nov., sp. nov. (Archosauria: Pterosauria) from the Romualdo Formation (Aptian/Albian), is herein described. The specimen (MPSC R 2357) was found at Sítio São Gonçalo, Santana do Cariri city (State of Ceará, northeast Brazil) and consists of the skull, atlas and axis, and represents one of the largest skulls of the Anhangueridae from the Araripe Basin described. The autapomorphies of the new pterosaur include the following characters: a premaxillary sagittal crest that is relatively long and high, beginning at the anterior part of the skull (rostrum) and extending to the 22nd pair of alveoli, not covering the nasoantorbital fenestra or the choanaes, and also the presence of 35 pairs of alveoli; smooth palatal ridge, which starts on the 5th pair of alveoli and ends on the 13th pair; palate is convex shaped in the anterior region; choanae not extending laterally; small and convex palatal elevation; the 5th, 6th and 7th alveoli smaller than the 4th and 8th; the alveoli decreasing in size from the 9th to the 12th and increasing from the 13th to 18th, and from the 18th to the 35th they are arranged in triplets. Furthermore, the lateral surface of the premaxillary crest shows grooves and tridimensional structures that may have housed blood vessels.
A three-dimensional and almost complete pterosaur mandible from the Crato Formation (Early Cretaceous of Northeastern Brazil), Araripe Basin, is described as a new species of a tapejarine tapejarid. Tapejarines are a particular group of toothless pterosaurs, characterized by well-developed cranial crests, downturned rostra, and have been proposed to represent frugivorous flying reptiles. Though comparatively well represented and distributed, the evolutionary history of the group is still poorly known, and the internal relationships of its members are not well understood. The new species here reported, named Aymberedactylus cearensis gen. et sp. nov., adds new data concerning the evolution of the group, concerning their morphology and geographical origin. It differs from known tapejarids due to its unusually elongate retroarticular process and a shallow fossa on the splenial exhibiting distinctive rugose texture. Furthermore, it exhibits a suite of basal and derived conditions within the Tapejaridae, demonstrating how their morphological traits probably evolved and that these forms were even more diverse than already acknowledged. The discovery of Aymberedactylus cearensis sheds new light on the evolutionary history of the Tapejarinae.
Pterosaurs are extinct flying reptiles, the first vertebrates to achieve powered flight. Our understanding of the evolutionary transition between basal, predominantly long-tailed forms to derived short-tailed pterodactyloids remained poor until the discovery of Wukongopterus and Darwinopterus in western Liaoning, China. In this paper we report on a new genus and species, Douzhanopterus zhengi, that has a reduced tail, 173% the length of the humerus, and a reduced fifth pedal digit, whose first phalange is ca. 20% the length of metatarsal III, both unique characters to Monofenestra. The morphological comparisons and phylogenetic analysis presented in this paper demonstrate that Douzhanopterus is the sister group to the ‘Painten pro-pterodactyloid’ and the Pterodactyloidea, reducing the evolutionary gap between long- and short-tailed pterosaurs.
The Tapejaridae is a group of unusual toothless pterosaurs characterized by bizarre cranial crests. From a paleoecological point of view, frugivorous feeding habits have often been suggested for one of its included clades, the Tapejarinae. So far, the presence of these intriguing flying reptiles has been unambiguously documented from Early Cretaceous sites in China and Brazil, where pterosaur fossils are less rare and fragmentary than in similarly-aged European strata.
In the Early Cretaceous Jehol Biota, the toothless pterosaurs flourished with the chaoyangopterids and tapejarids playing a key role in understanding the early diversity and evolution of the Azhdarchoidea. Unlike the more diverse tapejarids, the rarer chaoyangopterids are characterized by a long and low rostrum, supporting a close relationship with the huge azhdarchids. Unfortunately, our knowledge is still limited in the osteology, paleoecology, and taxonomy of the Chaoyangopteridae. As one of the best preserved skeletons, the type and only specimen of Jidapterus edentus provides an opportunity to understand the morphology and paleoecology of the chaoyangopterids.