This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs went extinct millions of years before humans evolved, humans have coexisted with pterosaur fossils for millennia. Before the development of paleontology as a formal science, these remains would have been interpreted through a mythological lens. Myths about thunderbirds told by the Native Americans of the modern Western United States may have been influenced by observations of Pteranodon fossils. These thunderbirds were said to have warred with water monsters, which agrees well with the co-occurrence of Pteranodon and the ancient marine reptiles of the seaway over which it flew. [1]
The formal study of pterosaurs began in the late 18th century when naturalist Cosimo Alessandro Collini of Mannheim, Germany published a description of an unusual animal with long arms, each bearing an elongated finger. He recognized that this long finger could support a membrane like that of a bat wing, but because the unnamed creature was found in deposits that preserve marine life he concluded that these strange arms were used as flippers. [2] The creature was restudied again in the very early 19th century by French anatomist Georges Cuvier, who recognized both that the creature was a reptile and that its "flippers" were wings. He called the creature the Ptero-dactyle, a name since revised to Pterodactylus . [3]
Although Cuvier's interpretation later became the consensus, it was just one of many early interpretations of the creature and its relatives, including that they were bats, strange birds, or the primordial handiwork of Satan himself. [4] Similar animals like the long-tailed Rhamphorhynchus and Gnathosaurus were soon discovered around Europe and it became obvious that earth was once home to a diverse group of flying reptiles. [5] The British anatomist Sir Richard Owen dubbed this vanished order the Pterosauria. Soon after, he described Britain's own first pterosaur, Dimorphodon . [6] Later in the 19th century pterosaurs were discovered in North America as well, the first of which was a spectacular animal named Pteranodon by paleontologist Othniel Charles Marsh. [7]
Various aspects of pterosaur biology invited controversy from the beginning. Samuel Thomas von Soemmering ignited a multi-century debate over how pterosaurs walked on the ground by suggesting they crawled on all fours like bats. August Quenstedt, by contrast, argued that they walked on their hind limbs. [8] In the early 20th century, Hankin and Watson in the first major study of pterosaur flight biomechanics concluded that on the ground these reptiles were altogether helpless and could only scoot along on their stomachs like penguins. [9] The debate gained steam in 1957 when William Stokes reported unusual tracks left by a four-footed animal he suspected was a pterosaur walking along the ground. [10] In 1984, Kevin Padian, who had recently argued that pterosaurs walked on their hind legs, dismissed Stokes's tracks as those of a crocodilian. [11] However, in the mid-1990s, Jean-Michel Mazin and others reported that fossil footprints in Crayssac, France were similar to those reported by Stokes from the US. Mazin's tracks were more obviously pterosaurian in origin and settled the debate in favor of pterosaurs walking on all fours. [10]
Pterosaur paleontology continues to progress into the 21st century. In fact, according to David Hone the early 21st century has seen more progress in pterosaur paleontology than in "the preceding two centuries" combined. He compared this transformative period in pterosaur paleontology to the Dinosaur Renaissance of the 1970s. [12] He also observed that roughly one-third of known pterosaurs were discovered during this brief interval. [13] One of the most notable of these was Darwinopterus , whose body resembled the more primitive long-tailed "rhamphorynchoids", while its skull resembled those of the more advanced short-tailed pterodactyloids. [14] These traits establish the species as an important transitional form, documenting one of the most important phases of pterosaur evolution. [15] Another important new species is Faxinalipterus minima , which might well be the world's oldest pterosaur. [16] The first confirmed pterosaur eggs were also reported from China during the early 21st century. [17]
The Cheyenne people of Nebraska believed in mythical thunderbirds and water monsters that were in endless conflict with each other. The thunderbirds were said to resemble giant eagles and killed both people and animals with arrows made of lightning. People occasionally discovered stony arrowheads thought to come from the thunderbirds' arrows. According to folklorist Adrienne Mayor, these supposed arrowheads were likely fossil belemnites, which were compared to missiles by other indigenous American cultures, like the Zuni people. [18]
The fossils of the Niobrara chalk may have been influential on these stories. The pterosaur Pteranodon and marine reptiles like mosasaurs are preserved in Niobrara Chalk deposits and associated remains may have been interpreted as evidence for antagonism between immense flying animals and serpentine aquatic reptiles. Fossils of the large toothed diving bird Hesperornis are also found in the Niobrara chalk, sometimes preserved inside specimens of large predatory marine reptiles. Observations of similar fossils in the past may have been seen as further evidence for thunderbird-water monster conflict. [19]
c. 1825
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In this paper Williston also described a new, relatively complete Nyctosaurus specimen. He noted that the only published trait distinguishing the genus from Pterodactylus was an absence of teeth and recommended synonymizing these two genera if "Nyctosaurus" teeth were ever found. [57]
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Pterodactylus is a genus of extinct pterosaurs. It is thought to contain only a single species, Pterodactylus antiquus, which was the first pterosaur to be named and identified as a flying reptile and one of the first prehistoric reptiles to ever be discovered.
Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous. Pterosaurs are the earliest vertebrates known to have evolved powered flight. Their wings were formed by a membrane of skin, muscle, and other tissues stretching from the ankles to a dramatically lengthened fourth finger.
Pteranodon ; from Ancient Greek πτερόν and ἀνόδων is a genus of pterosaur that included some of the largest known flying reptiles, with P. longiceps having a wingspan of over 6 m (20 ft). They lived during the late Cretaceous geological period of North America in present-day Kansas, Nebraska, Wyoming, South Dakota and Alabama. More fossil specimens of Pteranodon have been found than any other pterosaur, with about 1,200 specimens known to science, many of them well preserved with nearly complete skulls and articulated skeletons. It was an important part of the animal community in the Western Interior Seaway.
Tapejara is a genus of Brazilian pterosaur from the Cretaceous Period. Tapejara crests consisted of a semicircular crest over the snout, and a bony prong which extended back behind the head. It was a small pterosaur, with a wingspan of approximately 1.23–1.3 metres (4.0–4.3 ft).
Cearadactylus is a genus of large anhanguerid pterodactyloid pterosaur from the Romualdo Formation of Brazil, South America. Fossil remains of Cearadactylus dated back to the Albian stage of the Early Cretaceous period, about 112 million years ago. The only known species is C. atrox, described and named in 1985 by Giuseppe Leonardi and Guido Borgomanero. The name refers to the Brazilian state Ceará, and combines this with Greek daktylos, "finger", a reference to the wing finger of pterosaurs. The Latin atrox means "frightful", a reference to the fearsome dentition of the species.
Ornithocheirus is a pterosaur genus known from fragmentary fossil remains uncovered from sediments in the United Kingdom and possibly Morocco.
This phylogeny of pterosaurs entails the various phylogenetic trees used to classify pterosaurs throughout the years and varying views of these animals. Pterosaur phylogeny is currently highly contested and several hypotheses are presented below.
Germanodactylus is a genus of germanodactylid pterodactyloid pterosaur from Upper Jurassic-age rocks of Germany, including the Solnhofen Limestone. Its specimens were long thought to pertain to Pterodactylus. The head crest of Germanodactylus is a distinctive feature.
Anhanguera is a genus of pterodactyloid pterosaur known from the Early Cretaceous Romualdo Formation of Brazil and the Late Cretaceous Kem Kem Group of Morocco. This pterosaur is closely related to Ornithocheirus, but belongs in the family Anhangueridae. The generic name comes from the Tupi words añanga, meaning "spirit protector of the animals" + wera "bygone".
Ornithostoma is a genus of pterodactyloid pterosaur that lived during the Early Cretaceous period of Europe, around 110 million years ago. Ornithostoma was once thought to have been a senior synonym of the pteranodontid Pteranodon due to its toothless anatomy and prior naming.
Coloborhynchus is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. Coloborhynchus is known from the Lower Cretaceous of England, and depending on which species are included, possibly the Albian and Cenomanian ages as well. Coloborhynchus was once thought to be the largest known toothed pterosaur, however, a specimen of the closely related Tropeognathus is now thought to have had a larger wingspan.
Lonchodectes was a genus of lonchodectid pterosaur from several formations dating to the Turonian of England, mostly in the area around Kent. The species belonging to it had been assigned to Ornithocheirus until David Unwin's work of the 1990s and 2000s. Several potential species are known; most are based on scrappy remains, and have gone through several other generic assignments. The genus is part of the complex taxonomy issues surrounding Early Cretaceous pterosaurs from Brazil and England, such as Amblydectes, Anhanguera, Coloborhynchus, and Ornithocheirus.
Pteranodontoidea is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous of Asia, Africa, Europe, North America and South America. It was named by Alexander Wilhelm Armin Kellner in 1996. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Anhanguera, Pteranodon and all its descendants. The clade Ornithocheiroidea is sometimes considered to be the senior synonym of Pteranodontoidea, however it depends on its definition. Brian Andres in his analyses, converts Ornithocheiroidea using the definition of Kellner (2003) to avoid this synonymy.
Ardeadactylus is an extinct genus of ctenochasmatoid pterosaur known from the Late Jurassic Solnhofen limestone of Bavaria, southern Germany. It contains a single species, Ardeadactylus longicollum, which was originally thought to be a species of Pterodactylus, as P. longicollum.
Lonchodraco is a genus of lonchodraconid pterodactyloid pterosaur from the Late Cretaceous of southern England. The genus includes species that were previously assigned to other genera.
Aerodactylus is a pterosaur genus containing a single species, Aerodactylus scolopaciceps, previously regarded as a species of Pterodactylus.
Ornithocheiromorpha is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 92.5 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica, though most genera were recovered in Europe, Asia and South America. They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by pteranodontians and azhdarchoids. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus, but not Pteranodon.
Serradraco is a genus of Early Cretaceous pterodactyloid pterosaur from the Valanginian aged Tunbridge Wells Sand Formation in England. Named by Rigal et al. in 2018 with the description of a second specimen, it contains a single species, S. sagittirostris, which was formerly considered a species of Lonchodectes, L. sagittirostris. In 2020, Averianov suggested it did not belong in Lonchodectidae.
Wellnhopterus is an azhdarchid pterosaur recovered from the Late Cretaceous Javelina Formation in Texas that was previously identified as a thalassodromine. It consists of a set of upper and lower jaws, as well as some cervical vertebrae and a fragmentary long bone. In July 2021, the jaws were given the genus name "Javelinadactylus", with the type and only species as "J. sagebieli"; however, this article has now been retracted. In a paper published in December 2021, the complete holotype was independently named Wellnhopterus, with the only species being W. brevirostris. As of 2022, this is the formal name of this pterosaur.