Propterodactylus | |
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Holotype specimen | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Clade: | † Pterodactyliformes |
Clade: | † Caelidracones (?) |
Suborder: | † Pterodactyloidea (?) |
Genus: | † Propterodactylus Spindler, 2024 |
Species: | †P. frankerlae |
Binomial name | |
†Propterodactylus frankerlae Spindler, 2024 | |
Propterodactylus (meaning "before Pterodactylus ") is an extinct genus of transitional monofenestratan pterosaurs from the Late Jurassic Painten Formation of Germany. The genus contains a single species, P. frankerlae, known from a complete articulated skeleton. Before its naming, Propterodactylus was referred to as the "Painten pro-pterodactyloid" in the scientific literature.
The Propterodactylus holotype specimen, DMA-JP-2011/006, was discovered in April 2011 in the Rygol lime works quarry of the Painten Formation (Ulmense subzone) near Painten in Bavaria, Germany. The nearly-perfect specimen consists of an articulated complete skeleton with preserved soft tissues. [1] [2]
In 2024, Frederik Spindler described Propterodactylus frankerlae as a new genus and species of monofenestratan pterosaur based on these fossil remains. The generic name, Propterodactylus, combines the Ancient Greek suffix "προ-" ("pro-"), meaning "before", with the genus name Pterodactylus (derived from Greek words meaning "wing digit"). As such, the name references its position as a "forerunner" to the Pterodactyloidea, as well as its long-standing nickname as the "Painten pro-pterodactyloid". The specific name, frankerlae, honours Petra Hahn, née Frankerl, the deceased wife of Stephan Hahn, the discoverer of the Propterodactylus holotype. [1]
The holotype specimen of Propterodactylus belongs to either a juvenile or subadult individual. As such, it had not reached full skeletal maturity. The lack of fusion in several skeletal elements, such as the synsacrum, scapulocoracoid, carpals, and ischiopubic plate, indicate that the animal was not fully grown. However, the general robusticity of the bones and the presence of large carpals and tarsals indicate that it was also not a young juvenile. The total length of the holotype skull is 9.3 centimetres (3.7 in), and it has a wingspan of approximately 55 centimetres (22 in). [1]
Despite the 'transitional' nature of the Propterodactylus holotype, the skull has several features otherwise seen only in later diverging ctenochasmatoids, such as a nearly-circular orbit, a round posterior region of the skull, and generally horizontal occipital region. In general, the skull is most proportionally similar to Pterodactylus . [3] [1]
Based on the mosaic of features observed in the Propterodactylus holotype, it can be confidently regarded as transitional between the more basal monofenestratans such as darwinopterans and the later diverging lophocratian pterodactyloids. Several phylogenetic analyses have been conducted that include Propterodactylus (as the "Painten pro-pterodactyloid"), recovering generally similar results. Most recent studies have found Propterodactylus within the Pterodactyloidea, rather than the sister taxon to it. The results of Andres (2021) are displayed in the cladogram below, with Propterodactylus as the sister taxon to the Lophocratia within Pterodactyloidea, diverging after the fragmentary Kryptodrakon . [4] Pêgas (2024) independently recovered the same relationships, but with anurognathids outside of the Monofenestrata. [5] Similarly, Dalla Vecchia (2022) found Propterodactylus to be the basalmost pterodactyloid in an analysis with a smaller sample size, with the Darwinoptera as the sister taxon to Changchengopterus + Pterodactyloidea within the Monofenestrata. [6]
Some older studies, such as Vidovic & Martill (2018), discussed the Propterodactylus fossil material as belonging to a non-pterodactyloid immediately basal to the clade's divergence. [7] Similarly, Wang et al. (2017) recovered Propterodactylus as the sister taxon to the Pterodactyloidea, with Douzhanopterus as the sister to this clade. They further noted morphological similarities between Douzhanopterus and Propterodactylus indicating the "intermediate" status of both taxa. [8]
Caulkicephalus is a genus of anhanguerid pterosaur from the Isle of Wight off the coast of England. It lived during the Early Cretaceous period, about 130 million years ago.
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.
Tropeognathus is a genus of large pterosaurs from the late Early Cretaceous of South America. This genus is considered to be a member of the family Anhangueridae, however, several studies have also recovered it within another family called Ornithocheiridae. Both of these families are diverse groups of pterosaurs known for their keel-tipped snouts and large size. Tropeognathus is regarded as the largest pterosaur found in the Southern Hemisphere, only rivaled by the huge azhdarchids. The type and only species is Tropeognathus mesembrinus. Fossil remains of Tropeognathus have been recovered from the Romualdo Formation, which is a Lagerstätte located in the Santana Group of the Araripe Basin in northeastern Brazil.
Anurognathidae is a family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Five genera are known: Anurognathus, from the Late Jurassic of Germany; Jeholopterus, from the Middle to Late Jurassic of China; Dendrorhynchoides, from the Middle Jurassic of China; Batrachognathus, from the Late Jurassic of Kazakhstan; and Vesperopterylus, from the Early Cretaceous of China. Bennett (2007) suggested that the holotype of Mesadactylus, BYU 2024, a synsacrum, belonged to an anurognathid, though this affinity has been questioned by other authors. Mesadactylus is from the Late Jurassic Morrison Formation of the United States. Indeterminate anurognathid remains have also been reported from the Middle Jurassic Bakhar Svita of Mongolia and the Early Cretaceous of North Korea.
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.
Pterodactyloidea is one of the two traditional suborders of pterosaurs, and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal rhamphorhynchoids by their short tails and long wing metacarpals. The most advanced forms also lack teeth, and by the late Cretaceous, all known pterodactyloids were toothless. Many species had well-developed crests on the skull, a form of display taken to extremes in giant-crested forms like Nyctosaurus and Tupandactylus. Pterodactyloids were the last surviving pterosaurs when the order became extinct at the end of the Cretaceous Period, together with the non-avian dinosaurs and most marine reptiles.
Gallodactylidae is a group of pterosaurs within the suborder Pterodactyloidea. Gallodactylids differed from other related pterosaurs in several distinct features, including fewer than 50 teeth present only in the jaw tips, and rounded crests present on the rear portion of the skull and jaws but not near the ends of their snouts. At least some species possessed jaw flanges, possibly used to bissect hard-shelled prey.
Diopecephalus is a genus of pterodactyloid pterosaur from the Lower Tithonian of the Lithographic Limestone, Bavaria, Germany. The type and only species is D. kochi, although the name has been applied to Pterodactylus longicollum, with longicollum erroneously listed as the type species.
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.
Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. The Azhdarchoidea has been recovered as either closely related to the Ctenochasmatoidea, as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea, which in turn was also within the Ornithocheiroidea.
Uktenadactylus is a genus of anhanguerid pterodactyloid pterosaurs from the Lower Cretaceous Paw Paw Formation of Texas, United States and the Wessex Formation on the Isle of Wight, England. Fossil remains of Uktenadactylus dated back to the Early Cretaceous period, from about 125 to 100 million years ago.
Monofenestrata is an unranked group of pterosaurs that includes the family Wukongopteridae and the suborder Pterodactyloidea.
Euctenochasmatia is an extinct group of pterodactyloid pterosaurs. It was named by David Unwin in 2003 as the group that contains the most recent common ancestor of Pterodactylus and Ctenochasma, and all their descendants.
Archaeopterodactyloidea is an extinct clade of pterodactyloid pterosaurs that lived from the middle Late Jurassic to the latest Early Cretaceous periods of Africa, Asia, Europe and North America. It was named by Alexander Wilhelm Armin Kellner in 1996 as the group that contains Germanodactylus, Pterodactylus, the Ctenochasmatidae and the Gallodactylidae. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Pterodactylus, Ctenochasma and Gallodactylus and all its descendants. Although phylogenetic analyses that based on David Unwin's 2003 analysis do not recover monophyletic Archaeopterodactyloidea, phylogenetic analyses that based on Kellner's analyses, or the analyses of Brian Andres recover monophyletic Archaeopterodactyloidea at the base of the Pterodactyloidea.
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.
Kryptodrakon is an extinct genus of pterodactyloid pterosaur from the Middle to Late Jurassic with an age of approximately 162.7 million years. It is known from a single type species, Kryptodrakon progenitor. The age of its fossil remains made Kryptodrakon the basalmost and oldest pterodactyloid known to date.
Douzhanopterus is an extinct genus of monofenestratan pterosaur from the Late Jurassic of Liaoning, China. It contains a single species, D. zhengi, named by Wang et al. in 2017. In many respects, it represents a transitional form between basal pterosaurs and the more specialized pterodactyloids; for instance, its tail is intermediate in length, still being about twice the length of the femur but relatively shorter compared to that of the more basal Wukongopteridae. Other intermediate traits include the relative lengths of the neck vertebrae and the retention of two, albeit reduced, phalanx bones in the fifth digit of the foot. Phylogenetically, Douzhanopterus is nested between the wukongopterids and Propterodactylus, which is similar to Douzhanopterus in many respects but approaches pterodactyloids more closely elsewhere.