Recent discoveries of non-pterodactyloid pterosaurs in the Late Jurassic lacustrine deposits of Inner Mongolia, Hebei and Liaoning, northeastern China, represent a new assemblage in the Jurassic terrestrial ecosystem. Here, a new pterosaur skeleton is described from the Late Jurassic Tiaojishan Formation of Jianchang, western Liaoning, China, as Changchengopterus pani Lü, 2009 by the elongated cervical vertebrae, similar proportions of the wing fingers, metacarpals I-III disparate in length, and extremely-elongated zygapophyses. This discovery provides new information of the postcranial skeleton in the understanding of bony anatomy of Changchengopterus, and also further highlights its similarity with the recently-erected Darwinopterus and Wukongopterus. The three co-existing taxa are distinguished here by shape differences in the pedal digit V-2, but final confirmation requires discovery of more complete material. The new find expands the geographic range of Changchengopterus in neighbouring areas of Hebei and Liaoning.
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... Since the discovery of the Wukongopteridae and introduction of Monofenestrata for their sister group relationship with more derived pterosaurs (Wang et al., 2009;Lü et al., 2010) it is known that the pterodactyloid transition was modular, with the derived skull pattern evolving earlier and seemingly independent from postcranial modifications. Thus, along with fragmentary mosaic forms and further discoveries (Zhou and Schoch, 2011;Martill and Etches, 2013;citations in Tischlinger and Frey, 2013;Codorniú et al., 2016;Wang et al., 2017;Zhou et al., 2021), the initial steps towards the pterodactyl transition could be clarified to a certain extent. ...
... The anterior tip is widened to form a strengthened contact with the other hyoid. This trait is also seen in Darwinopterus , Changchengopterus (Zhou and Schoch, 2011), Gallodactlyus (Fabre, 1976, figure 2, if not preserving the basibranchial, see Jiang et al., 2020), as well as a specimen from Painten assigned to Pterodactylus (Augustin et al., 2022). ...
... For anurognathids, cervical ribs are stated, but not well-documented (Dalla Vecchia, 2002;Bennett, 2007; but see Lü et al., 2018), leading Hone (2020) to evaluate them as reduced or absent. In the more primitive Changchengopterus cervical ribs are apparently absent (Zhou and Schoch, 2011). Dorsal vertebrae. ...
... Among pterosaurs a short, deep, relatively broad-based cristospine, comparable to that present in NHMUK PV R37110, seems to be restricted to darwinopterans and is observed, for example, in Darwinopterus (e.g., Lü et al., , 2011Wang et al., 2010), Kunpengopterus , and, seemingly, in Changchengopterus (Zhou & Schoch, 2011). The sternal cristospine of Fenghuangopterus also appears to be broad-based , but the cristospine is relatively more elongate than in darwinopterans and its apparent robustness may be partly due to crushing. ...
... The lateral apex of the fibula is more rounded and the shaft, which is closely appressed to the tibia with no interosseous space, is relatively robust and more than one-half the width of the tibia shaft. The preserved portion of the tibiafibula of Ceoptera corresponds closely in all anatomical details to those of other darwinopterans, including Darwinopterus (Lü et al., 2011; HGM 41HIII-0309A), Changchengopterus (Zhou & Schoch, 2011; PMOL AP-00010), and Wukongopterus (Wang et al., 2009;IVPP V15113). ...
... Their spatial orientation within the wing membrane would have been variable depending on the respective wing position. In general, however, they were arranged approximately vertically relative to the bones of the forelimb and parallel to subparallel relative to the wing finger (Wellnhofer, 1987;Chatterjee and Templin, 2004;Kellner et al., 2010;Zhou and Schoch, 2011;Hone et al., 2015). ...
... Notably, the Daxishan locality is a major site where significant vertebrate fossils of the Middle-Late Jurassic Yanliao Biota (159-166 Ma) have been recovered (Sullivan et al., 2014;Xu et al., 2016;Zhou and Wang, 2017). To date, this locality has yielded fossil representatives of several vertebrate groups, including early mammals (Luo et al., 2011;Bi et al., 2014;Meng et al., 2017), feathered dinosaurs Zheng et al., 2009;Li et al., 2010;Xu et al., 2011), pterosaurs (e.g., Lü et al., 2011Zhou and Schoch, 2011;Cheng et al., 2012;Zhou, 2014;Wang et al., 2017;, and undescribed turtles and palaeoniscoid fishes (Gao et al., 2015). In addition to the new hynobiid-like salamander described herein, specimens of the cryptobranchoid salamander Chunerpeton tianyiensis and other undescribed caudates were reported from this same fossil locality (Sullivan et al., 2014), albeit none of these materials have yet been described. ...
Hynobiids are a group of small- to moderate-sized salamanders living primarily in Asia. They are a primitive crown-group clade, with a poor fossil record. Several hynobiid-like taxa have been discovered from the Lower Cretaceous strata of northern China during the last 20 years, with Liaoxitriton zhongjiani and Nuominerpeton aquilonaris identified as the oldest known stem hynobiids. However, the record of pre-Cretaceous hynobiid-like taxa is only known by Liaoxitriton daohugouensis, of which both the morphology and the congeneric status with L. zhongjiani remain problematic. Here, we report on a new hynobiid-like salamander, Linglongtriton daxishanensis, gen. et sp. nov., on the basis of two specimens from the Upper Jurassic Lanqi/Tiaojishan Formation (∼160 Ma) of Liaoning Province, China. Linglongtriton is diagnosed by a unique combination of features revealed by both observation under microscope and micro-computed tomography (μCT) scan of the holotype, including nasals separated from each other at the midline; prootic, opisthotic, and exoccipital retained as discrete elements; dentary with a lateral groove; articular not ossified; metacarpal III enlarged; a single centrale; and distal tarsals 4 and 5 fused into a single element. Phylogenetic analysis identified Linglongtriton and several other hynobiid-like taxa, including Liaoxitriton daohugouensis, as stem hynobiids, thereby extending the temporal range of the stem by at least 40 Ma—from the Early Cretaceous (Aptian–Barremian) to the Middle Jurassic (Bathonian). Comparative study of Linglongtriton with living and fossil hynobiids sheds new lights on the evolution and developmental mechanisms of several characters, including nasal separation and tarsal elements.
... Whereas CVA is more limited in its taxonomic information due to taphonomy, the three-dimensional preservation of CVB allows for a more detailed comparison. Howse (1986) considered the possession of cervical ribs as a characteristic of basal pterosaurs, but reduced cervical ribs are known from several pterodactyloid clades (Zhou and Schoch 2011), including the highly derived Azhdarchidae (Witton and Naish 2008). Cervical ribs are not preserved in the holotype of the non-pterodactyloid monofenestratan Darwinopterus modularis Lü, Unwin, Jin, Liu, and Ji, 2010, but there appear to be anterolateral projections on several vertebrae which may correspond to cervical rib facets. ...
The current understanding of UK Middle Jurassic pterosaur taxonomy is under-developed, leading to it being previously considered a time of low diversity. This is despite the presence of a productive but under-studied pterosaur-bearing horizon extending over parts of Oxfordshire and Gloucestershire. This unit, informally called the Stonesfield Slate, is part of the Great Oolite Group and it produces the largest number of Middle Jurassic pterosaurs. There are over 200 specimens distributed across museums in the United Kingdom, America, and Australia, almost all of which are accessioned under the genus Rhamphocephalus and referred to three species: the type species Rhamphocephalus prestwichi, Rhamphocephalus bucklandi, and Rhamphocephalus depressirostris. This study reviews the British Middle Jurassic Pterosauria assemblage, evaluating both their systematics and taxonomic diversity. The holotype of Rhamphocephalus, an isolated skull table, is found to be a misidentified crocodylomorph skull and the genus is considered a nomen dubium. The holotype of Rhamphocephalus bucklandi is identified as missing and that of Rhamphocephalus depressirostris has characters diagnostic at a family level, not a generic or specific one. Both species are considered dubious. Detailed examination of the entire assemblage shows that rather than being monogeneric, the assemblage contains at least five pterosaur taxa, representing three families. This diversity includes the potential earliest occurrences of both Monofenestrata and Pterodactyloidea. A new genus, Klobiodon rochei gen. et sp. nov. is described based on a well-preserved mandible. The English Bathonian pterosaur assemblage is shown to be diverse and indicates that, as has been suggested in other studies, the low-diversity signal in the Middle Jurassic is at least partially artificial.
... So far, six specimens of this non-pterodactyloid clade have been described and referred to three genera and five species ( Wang et al., 2009;Wang et al., 2010;Lü et al., 2010;Lü et al., 2011a). In addition, there is Changchengopterus pani, known from two individuals (Lü, 2009;Zhou & Schoch, 2011), that is regarded as a potential wukongopterid ( Wang et al., 2009;Cheng et al., 2017), and another three undetermined specimens (Lü et al., 2011b;Cheng et al., 2016;Cheng et al., 2017). Furthermore, there are several undescribed fossils scattered throughout many collections in China that most certainly belong to this clade. ...
The Wukongopteridae compose a non-pterodactyloid clade of pterosaurs that are the most abundant flying reptiles in the deposits of the Middle-Late Jurassic Yanliao Biota. Until now, five species of three genera and two additional unnamed specimens have been described. Here we report on a new material, IVPP V 23674, that can be referred to the wukongopterid Kunpengopterus sinensis due to several features such as a comparably short nasoantorbital fenestra, the dorsally rising posterodorsal margin of the ischium, and the very short first pedal phalanx of digit V relative to metatarsal IV. IVPP V 23674 provides the first view of a wukongopterid palate, which differs from all other pterosaurs by having a very large postpalatine fenestra and laterally compressed choanae, indicating that the evolution of the pterosaur palate was more complex than previously thought. Sesamoid bones at the dorsal side of manual unguals are present and are reported for the first time in a wukongopterid suggesting an arboreal life-style for these pterosaurs.
Pterosaurs were the first powered flying vertebrates, with a fossil record that stretches back to about 230 million years before present. Most species are only known from one to three specimens, which are most often fragmentary. However, Rhamphorhynchus muensteri is known from numerous excellent specimens, including multiple specimens with soft tissue preservation. As such, Rhamphorhynchus muensteri is one of the only pterosaurs amenable to analysis for intraspecific variation. It has been previously predicted that elements directly involved in the flight apparatus, such as those of the forelimb, will be more highly constrained in their proportions than other parts of the skeleton. We investigated the degree of variation seen in elements and body parts of Rhamphorhynchus , which represents the best model system among pterosaurs for testing these expectations of intraspecific variation. We recover evidence for high levels of constraint throughout the appendicular and axial elements (head, neck, torso, tail, forelimbs, hindlimbs), suggesting that all were important for flight. We further find that tail variation increases among the largest specimens, suggesting reduced constraint and/or stronger sexual selection on the tail in more mature individuals.
As the lowest horizon of the Early Cretaceous Jehol Biota, the Huajiying Formation of the northern Hebei Province, China is rich with avians and feathered dinosaurs, but lacks pterosaur record. Here the first pterosaur fossil is reported from the Huajiying Formation. The new pterosaur specimen is characterized by an unusual pedal configuration of a short and spread metatarsus with elongate digits, showing a close resemblance to the dentulous Ornithocheiroidea. The pedal configuration is diverse in pterosaurs, and is often associated with ecological adaptations. In contrast to the general pattern of the elongate metatarsus and short digits, the elongate digits of the dentulous ornithocheiroids (e.g. boreopterids and istiodactylids) might offset the shortened metatarsus to enlarge the pedal surface for paddling, representing a new strategy in adaptation to the aquatic environment.
The anurognathids are peculiar pterosaurs characterized by broad skulls with very short rostra and broadly arched jaws. The presence of distinct or confluent external naris and antorbital fenestra in these pterosaurs has been debated in the last years. The relatively well-preserved specimens of Batrachognathus volans show that the antorbital fenestra was confluent with the orbit forming an enormous orbitoantorbital fenestra. This feature is evident also in Jeholopterus ningchengensis. The consequent modification of the matrices of two recently published phylogenetic analyses about the in-group pterosaur relationships shows that the Anurognathidae are a derived clade of non-monofenestratan pterosaurs. Anurognathidae (including also 'Dimorphodon' weintraubi according to the definition by Hone 2020) are still a scarcely known clade because only a few specimens have been adequately described in the literature.
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The first scientific description of a pterosaur was published in 1784 by Cosimo Alessandro Collini, a former secretary of Voltaire and at that time curator of the natural history cabinet of Karl Theodor, Elector of Palatinate and Bavaria. The specimen came from one of the main sources of such fossils, the Late Jurassic lithographic limestones of northern Bavaria, and Collini, after much deliberation, interpreted it as the skeleton of an unknown marine creature (Collini 1784).
In 1801, Georges Cuvier, on the basis of Collini’s description and figure, identified the mysterious animal as a flying reptile (Cuvier 1801), for which he later coined the name ‘Ptero-Dactyle’ (Cuvier 1809). Cuvier’s basically correct interpretation of the ‘winged finger’ marked the beginning of the study of pterosaurs as an extinct group of flying reptiles.
In the two centuries which have elapsed since those first efforts to understand what have been considered bizarre fossils, the study of pterosaurs has developed enormously. Some of the basic questions about them have long been solved: pterosaurs were neither birds, nor bats, as was suggested by various authors of the early nineteenth century, but a peculiar group of vertebrates which acquired the ability to fly in an original way, using a membrane attached to a single finger of the hand. From the few fossils from the Bavarian lithographic limestones known to Cuvier and his contemporaries, the number of pterosaur specimens has increased enormously, starting with the Early Jurassic specimens from Lyme Regis found by Mary Anning in the 1820s
Over 30 skeletons and dozens of isolated bones of the Liassic pterosaur Dorygnathus have been recovered from the Early Jurassic (Toarcian) of Baden-Württemberg and Lower Saxony in Germany, and from Nancy, France. All but one specimen have been assigned to the species D. banthensis; the exception was assigned to a larger species, D. 'mistelgauensis', which new discoveries suggest is simply a large individual of D. banthensis. The form of the lower jaw and premaxillary teeth are diagnostic for the genus, as are several other features. Here I review the history of the understanding of Dorygnathus, describe the known specimens in public repositories, and characterize the general morphology and systematic position of the genus. Dorygnathus is distinguished by its extremely large anterior teeth (four premaxillary and three or four anterior dentary teeth), which are proportionally larger than in any other pterosaur, Its deep maxilla gives the skull a high, straight, gradual slope, and its long, deepened, upwardly curved mandibular symphysis is diagnostic for the taxon. Other features such as the proportions of the wing elements, the form of the pelvis, and the shape and proportions of the toes are equally characteristic. Dorygnathus is most closely related to Rhamphorhynchus and the Pterodactyloidea, and represents this lineage in the Early Jurassic of Europe.
A cladistic analysis based on 39 terminal taxa and 74 characters (several multistate) using PAUP (Phylogenetic Analysis Using Parsimony) (3.1.1 for MacIntosh and 4.0b10 for Microsoft Windows) presents a new hypothesis of pterosaur inter-relationships. This study suggests that the most primitive taxon is the Anurognathidae, followed by Sordes and all remaining pterosaurs. Dendrorhynchoides is confirmed as a member of the Anurognathidae, being closely related to Batrachognathus. Preondactylus occupies a more derived position than Sordes , which questions its previous assignment as the most primitive pterosaur. The hypothesis of rhamphorhynchoid paraphyly is confirmed, with the Rhamphorhynchidae more closely related to the Pterodactyloidea than to more basal forms. The Pterodactyloidea shows a basal dichotomy: the Archaeopterodactyloidea and the Dsungaripteroidea. The Archaeopterodactyloidea is formed by Pterodactylus + Germanodactylus and a clade formed by Gallodactylidae + Ctenochasmatidae. The Nyctosauridae occupies the basal position within dsungaripteroids and is followed by the Pteranodontoidea and the Tapejaroidea. Pteranodontoids have Pteranodon at the base, followed stepwise by Istiodactylus, Ornithocheirus and the Anhangueridae. Tapejaroids are composed of the Dsungaripteridae at the base followed by the Tapejaridae and the Azhdarchidae.
Major trends within pterosaur evolutionary history are: general increase in size (wing span and body); increase of wing metacarpal and pteroid; decrease of proportional length of the second and third wing phalanx relative to the first; gradual increase of rostrum (anterior to external nares); and anterior shift of the skull-mandible articulation. Cranial crests are present in most pterodactyloids, but markedly in the Ornithocheiroidea, where all taxa show some sort of crest on the skull. The loss of teeth, previously assumed to have occurred independently in several lineages, seems to be a general trend among dsungaripteroids.
Several nodes recovered by this analysis are supported by very few characters, a result at least partially attributable to the limited available information from several taxa due to poor preservation and/or preparation.
Remains of pterosaurs, the dominant aerial vertebrate throughout much of the Mesozoic were, until relatively recently, almost exclusively known from marine and marginal marine sediments of western Europe and North America. Prior to the 1960s Mesozoic deposits in the former Soviet Union and Mongolia had produced very few pterosaurs, but, in the last thirty years, many remains, including some from continental environments, have been found. Most important among these are Sordes and Batrachognathus from the Late Jurassic of Karatau in Kazakhstan, Azhdarcho from the Late Cretaceous of the Kyzylkum desert in Uzbekistan, and a number of new Mongolian pterosaurs, including a possible anurognathid from the Mid Jurassic of Bakhar, a dsungaripterid from the early Early Cretaceous of Tatal and an ornithocheirid from the late Early Cretaceous of Khuren‐Dukh. Although already documented to some extent, the significance of these discoveries remains underappreciated.The discovery and collection of these pterosaurs is described and the bearing of some of the more important material on current problems of pterosaur biology is discussed. We confirm Sharov's observation that the hind limbs of Sordes form an integral part of the flight apparatus, attached externally to the cheiropatagium and internally to the uropatagium, which is supported and manipulated by the fifth toe. The former Soviet and Mongolian pterosaurs also help to fill a number of important morphological and temporal gaps in the pterosaur fossil record and provide the best available evidence of pterosaurs from continental environments. With the exception of insectivores, these and other continental pterosaurs appear to have pursued lifestyles similar to those of their marine counterparts, leading us to suspect that pterosaurs largely failed to exploit terrestrial habitats.
For more than 100 years, most pterosaur workers have accepted the traditional reconstruction of the pteroid, articulated in the fovea of the preaxial carpal and directed medially along the anterior margin of a small propatagium in order to control it. A few workers have rejected the traditional reconstruction and advocated an alternative reconstruction with the pteroid directed anteriorly and controlling a large propatagium extending laterally past the wrist. Articulated specimens of a wide range of pterosaurs preserve a sesamoid associated with the tendon of M. flexorcarpi ulnaris in the fovea of the preaxial carpal. No specimen preserves the pteroid articulated in the fovea, and because the sesamoid articulated there, the pteroid could not. Therefore, both the traditional and alternative reconstructions, which are based on the assumption that the pteroid articulated in the fovea, are falsified. The preaxial carpal acted as a strut to increase the leverage of M. flexor carpi ulnaris for wrist extension. The pteroid articulated with the side of the carpal, was directed medially toward the shoulder, and could be extended and depressed to control the propatagium.