When was the first Pteranodon fossil found?
Pteranodon
"Pteranodon" was the first pterosaur found outside of Europe. Its fossils first were found by Othniel Charles Marsh in 1870, in the Late Cretaceous Smoky Hill Chalk deposits of western Kansas. These chalk beds were deposited at the bottom of what was once the Western Interior Seaway, a large shallow sea over what now is the midsection of the North American continent. These first specimens, YPM 1160 and YPM 1161, consisted of partial wing bones, as well as a tooth from the prehistoric fish "Xiphactinus", which Marsh mistakenly believed to belong to this new pterosaur (all known pterosaurs up to that point had teeth). In 1871, Marsh named the find ""Pterodactylus oweni"", assigning it to the well-known (but much smaller) European genus "Pterodactylus". Marsh also collected more wing bones of the large pterosaur in 1871. Realizing that the name he had chosen had already been used for Harry Seeley's European pterosaur species "Pterodactylus oweni" in 1864, Marsh re-named his giant North American pterosaur Pterodactylus occidentalis, meaning "Western wing finger," in his 1872 description of the new specimen. He also named two additional species, based on size differences: Pterodactylus ingens (the largest specimen so far), and Pterodactylus velox (the smallest).
Timeline of troodontid research
This timeline of troodontid research is a chronological listing of events in the history of paleontology focused on the troodontids, a group of bird-like theropod dinosaurs including animals like "Troodon". Troodontid remains were among the first dinosaur fossils to be reported from North America after paleontologists began performing research on the continent, specifically the genus "Troodon" itself. Since the type specimen of this genus was only a tooth and "Troodon" teeth are unusually similar to those of the unrelated thick-headed pachycephalosaurs, "Troodon" and its relatives would be embroiled in taxonomic confusion for over a century. "Troodon" was finally recognized as distinct from the pachycephalosaurs by Phil Currie in 1987. By that time many other species now recognized as troodontid had been discovered but had been classified in the family Saurornithoididae. Since these families were the same but the Troodontidae named first, it carries scientific legitimacy.
Saurornithoides
Originally, only one or possibly two individuals of "Saurornithoides" were known, closely associated within the same layer of the Djadochta Formation of Mongolia. The fossils were found on 9 July 1923 by a Chinese employee of an American Museum of Natural History expedition, Chih. The material contained a single skull and jaw in association, and vertebrae, a partial pelvis, hindlimb and foot associated nearby. More bones were initially included but later shown to belong to "Protoceratops". Henry Fairfield Osborn at first intended to name the animal "Ornithoides", the "bird-like one", and in 1924 mentioned this name in a popular publication but without a description so that it remained an invalid "nomen nudum". He then formally described the remains in the same year, finding them to be a new genus and species, which he named "Saurornithoides mongoliensis". The generic name was chosen because of the bird-like bones of the taxon, which was thought to represent a megalosaurian, translating as "saurian with bird-like rostrum". "Saurornithoides" was noted to resemble "Velociraptor", although more sluggish according to Osborn. The holotype specimen is AMNH 6516. This specimen was the first troodontid skeleton found, though at the time the connection with "Troodon", then known only from its teeth, was not realised.
In 1964, another specimen was described from the Late Cretaceous of Mongolia. The first specimen ever collected by a professional Mongolian palaeontologist, it was given the specimen number IGM 100/1. In 1974, it was described by Rinchen Barsbold as a new species, "Saurornithoides junior". It was assigned to the genus based on cranial comparisons, as well as the similar provincialism. However, since the description, many more troodontids have been described with cranial material, and as such, a 2009 study on "Saurornithoides" reassigned this species to its own genus, "Zanabazar".
Paranthodon
In 1845, amateur geologists William Guybon Atherstone and Andrew Geddes Bain discovered several fossils near Dassieklip, Cape Province, in the Bushmans River Valley. This was the first dinosaur find in all of the Southern Hemisphere and Africa. In 1849 and 1853, Bain sent some of the fossils to palaeontologist Richard Owen for identification. Among them was an upper jaw Bain referred to as the "Cape "Iguanodon"", so the site was named "Iguanodonhoek". Atherstone published a short paper about the discovery in 1857, but lamented in 1871 that it had thus far received no attention in London. Only in 1876 were a series of specimens from the collection named by Owen as "Anthodon serrarius", basing the generic name on the resemblance of the teeth to a flower. The partial holotype skull BMNH 47337, the left jaw BMNH 47338, the matrix BMNH 47338 including bone fragments and impressions of the anterior skull, and the vertebrae BMNH 47337a were all assigned to "Anthodon." In 1882, Othniel Charles Marsh assigned "Anthodon" to Stegosauridae based on BMNH 47338, and in 1890, Richard Lydekker found that although "Anthodon" was a pareiasaur, its teeth were similar to those of the Stegosauridae. Lydekker in 1890 also corrected a mistake of Owen, who had incorrectly summarized all the material as coming from a single locality, whereas there was separate material from two clearly distinct localities.
Paleontology in Pennsylvania
One of the first major fossil finds of the 20th century in Pennsylvania was the 1902 discovery of dinosaur tracks at a fisher's Quarry near Graterford in Montgomery County. Another prominent early 1900s discovery was a lower jaw from the amphibian "Calamops" was discovered in the Late Triassic Stockton Formation at Holicong. This is the oldest vertebrate fossil from either the Hartford or Newark Basins. In 1923 the Reading Railroad tunnel at Gwynned was converted into an open cut, exposing many Late Triassic fossils. Later, in 1933, two "Anchisauripus" tracks were discovered near Yocumtown in York County. Dinosaur tracks dating back to the Late Triassic were discovered near the Gettysburg battle sites in 1933.
Pteranodon
Specimens assigned to "Pteranodon" have been found in both the Smoky Hill Chalk deposits of the Niobrara Formation, and the slightly younger Sharon Springs deposits of the Pierre Shale Formation. When "Pteranodon" was alive, this area was covered by a large inland sea, known as the Western Interior Seaway. Famous for fossils collected since 1870, these formations extend from as far south as Kansas in the United States to Manitoba in Canada. However, "Pteranodon" specimens (or any pterosaur specimens) have only been found in the southern half of the formation, in Kansas, Wyoming, and South Dakota. Despite the fact that numerous fossils have been found in the contemporary parts of the formation in Canada, no pterosaur specimens have ever been found there. This strongly suggests that the natural geographic range of "Pteranodon" covered only the southern part of the Niobrara, and that its habitat did not extend farther north than South Dakota.
Zářecká Lhota
In 1880, first and still the only known Czech pterosaur fossil (now known as "Cretornis hlavaci") was found in a nearby quarry.
Pteranodon
Some very fragmentary fossils belonging to pteranodontian pterosaurs, and possibly "Pteranodon" itself, have also been found on the Gulf Coast and East Coast of the United States. For example, some bone fragments from the Mooreville Formation of Alabama and the Merchantville Formation of Delaware may have come from "Pteranodon", though they are too incomplete to make a definite identification. Some remains from Japan have also been tentatively attributed to "Pteranodon", but their distance from its known Western Interior Seaway habitat makes this identification unlikely.
"Pteranodon longiceps" would have shared the sky with the giant-crested pterosaur "Nyctosaurus". Compared to "P. longiceps", which was a very common species, "Nyctosaurus" was rare, making up only 3% of pterosaur fossils from the formation. Also less common was the early toothed bird, "Ichthyornis".
Gannat
The most ancient discoveries in Gannat, ancestors of the rhinoceros from the end of the Oligocene and start of the Miocene, date back 23 million years. Gannat seems to have been a veritable cemetery for these creatures, so many of their remains have been discovered. Also found are fossils of fish, reptiles, tortoises, crocodiles, galliform birds, mammals, marsupials, insectivores, rodents and carnivores. The site is particularly rich in rhinoceros. From 1854, Duvernoy has even described a specimin as "Acerotherium gannatense" (the official name is "Diaceratherium lemanense"). The most complete rhinoceros skeleton was discovered in 1993 when significant fossils were uncovered at the Sichaux quarry by paleontologist François Escuillié (notable discoverer of a small proboscidian mammal, ancestor of the elephant from 50 million years ago). Escuillié was behind the establishment of the Rhinopolis Association in 1994 and founder of Eldonia, a society for those specialising in the renovation of fossils and whose activities are linked to Rhinopolis. The Rhinopolis Association is still active in the Gannat quarries and, since the 1990s, has discovered numerous rhinoceros bones.
Paleontology in Kansas
In 1870, O. C. Marsh led a paleontological expedition into the western United States on behalf of Yale University. Late in the November of that year they visited the area around Fort Wallace. They spent several days prospecting for fossils in modern Wallace and Logan Counties along the banks of the Smoky Hill River. Persevering against cold weather, they made several finds. One was the nearly complete skeleton of the mosasaur "Clidastes" that took four days to excavate. Among the fossils discovered by Marsh's crew in western Kansas were the far ends of two pterosaur wing metacarpals. These were the first scientifically documented fossils of the pterosaur that would later be named "Pteranodon". This formal naming occurred six years later, in 1876.
Palaeochenoides
Only a single species, Palaeochenoides mioceanus, is known to date. The first fossil assigned to it – a distal right femur piece – was found near the source of the Stono River in Charleston County, South Carolina (United States). At first it was believed to be from the Early Miocene Hawthorne Formation – its specific name, seemingly referring to the "Miocene ocean" as presumed habitat but actually a simple spelling error for "miocaenus", "from the Miocene", that was never corrected and hence became valid – alludes to this. But in fact no Hawthorne Formation rocks were known in the Charleston region when the fossil was found, and consequently modern authors consider a Chattian (Late Oligocene) age more likely and suggest the fossil came from the Cooper or Chandler Bridge Formation. Specimen MCZ 2514, a distal left tarsometatarsus fragment from the Ashley River, was more tentatively assigned to "P. mioceanus" later on; it was also erroneously believed to be from the Hawthorne Formation.
Pterygotus
Though early discoveries of "Pterygotus" were confined to England and North America, fossil finds throughout the 20th century would help establish that "Pterygotus" as a genus achieved a nearly cosmopolitan distribution. The first eurypterid to be discovered in Australia was "Pterygotus australis", whose fossils were found in the Ludlow age Melbourne Group of the Dargile Formation. The fossils referred to "P. australis", consisting of four fragments making up about half of a segment that were discovered during the process of excavations beneath Melbourne during the construction of new drainage works for the city in 1899. The fragmentary fossils closely resemble fossils of "Erettopterus bilobus" (classified as a species of "Pterygotus" at the time), which might make their assignment to "Pterygotus" questionable.
Protosphyraena
As is the case with many fossil vertebrates discovered by 19th century paleontologists, the taxonomy of "Protosphyraena" has had a confusing history. Fossil pectoral spines belonging to this taxon were first recognized in 1822, from chalk deposits in England, by Gideon Mantell, the physician and geologist who also discovered the dinosaur "Iguanodon". In 1857, the fish was named "Protosphyraena ferox" by the renowned American naturalist and paleontologist, Joseph Leidy, based on Mantell's English finds. Earlier, Leidy had published an illustration of a "Protosphyraena" tooth from the Cretaceous-aged Navesink Formation of New Jersey (Maastrichtian), but mistakenly identified is as having come from a dinosaur. During the 1870s, B. F. Mudge, a fossil collector supplying material to rival paleontologists Edward Drinker Cope and Othniel Charles Marsh, discovered a number of specimens of "Protosphyraena" in Niobrara exposures in Rooks and Ellis counties in Kansas and sent them back east. Between 1873 and 1877, Cope renamed three species based on Mudge's specimens, all of which would eventually be recognized as belonging to the genus "Protosphyraena": "Erisichte nitida", ""Portheus" gladius", and ""Pelecopterus" pernicciosus". Between 1895 and 1903, paleontologists in America and England, including Arthur Smith Woodward (1895), Loomis (1900), O. P. Hay (1903), in a series of important works, reviewed the genus, adding much to our understanding of this fish.
Pteranodontia
Pteranodontians were among the last of the world's pterosaur faunas. The species "Piksi barbarulna" and a few potential pteranodontids and nyctosaurs have all been found dating from the Campanian to the Maastrichtian ages of the Late Cretaceous period.
Pteranodon
Fragmentary fossils assigned to "Pteranodon" have also been discovered in Skåne, Sweden.
Pteranodon
A number of additional species of "Pteranodon" have been named since the 1870s, although most now are considered to be junior synonyms of two or three valid species. The best-supported is the type species, "P. longiceps", based on the well-preserved specimen including the first-known skull found by S. W. Williston. This individual had a wingspan of . Other valid species include the possibly larger "P. sternbergi", with a wingspan originally estimated at . "P. occidentalis", "P. velox", "P. umbrosus", "P. harpyia", and "P. comptus" are considered to be "nomina dubia" by Bennett (1994) and others who question their validity. All probably are synonymous with the more well-known species.
Martin Pickford
From 1971 to 1978 Pickford had carried out extensive research in the Tugen Hills under a permit issued by the Kenyan Office of the President. During the surveys Pickford and his team found many important fossils ranging in age from 15 million to 2 million years old. In 1974 he found the first hominid fossil from the 6-million-year-old Lukeino Formation (published in Nature in 1975), a lower molar, which is today included in the hypodigm of Orrorin tugenensis.
Pteranodon
"Pteranodon" fossils are known primarily from the Niobrara Formation of the central United States. Broadly defined, "Pteranodon" existed for more than four million years, during the late Coniacian to early Campanian stages of the Cretaceous period. The genus is present in most layers of the Niobrara Formation except for the upper two; in 2003, Kenneth Carpenter surveyed the distribution and dating of fossils in this formation, demonstrating that "Pteranodon sternbergi" existed there from 88 to 85 million years ago, while "P. longiceps" existed between 86 and 84.5 million years ago. A possible third species, which Kellner named "Geosternbergia maysei" in 2010, is known from the Sharon Springs member of the Pierre Shale Formation in Kansas, Wyoming, and South Dakota, dating to between 81.5 and 80.5 million years ago.
Stephen L. Brusatte
Brusatte has discovered more than a dozen new species of vertebrate fossils. His breakthrough in the study of dinosaur fossils was while at the University of Chicago with Paul Sereno. Having discovered the skull, jaw and neck fossils of a 95-million-year-old theropod in the Elrhaz Formation of Niger in 1997, Sereno was looking for a competent student to analyse it. Brusatte took the opportunity in 2004, completed the project in 2005, and published his findings in 2007 with Sereno. The animal was found to be a new species of "Carcharodontosaurus", which they named "C. iguidensis". He estimated that the complete skull would be more than five feet long, one of the biggest skulls of a known carnivorous dinosaur. This was followed by the description of another new theropod from the Elrhaz Formation in January 2008, "Kryptops palaios". Another significant discovery was from China in 2014. Alongside Chinese paleontologist Lü Junchang and others, Brusatte described a 66-million-year-old dinosaur, "Qianzhousaurus sinensis", which was closely related to the famous "T. rex". Due to its long snout, it was given the nickname "Pinocchio rex".
Pterasteridae
Fossil pterasterids have been found as early as the upper Campanian of the Cretaceous period.
Patranomodon
The skull fossil of "Patranomodon" was found in the "Eodicynodon" Assemblage Zone of South Africa, belonging to the lowest biozone of the Beaufort Group. The Beaufort Group time period extends from the middle of the Permian to the early Triassic era. It is one of the three main subdivisions of the Karoo Supergroup in what today is southern Africa. Rubidge and Hopson were the first to discover the skull of "Patranomodon". These paleobiologists also named "Patranomodon" and were the first to publish literature on it starting in 1990. The most abundant remains of "Patranomodon" were found on the Eastern Cape of South Africa; however, fossil parts were also found in Europe, China, as well as India, which indicated migration occurring among these terrestrial creatures. The paleontologist Nyaphuli collected the fossil of this creature in South Africa and gave it the species name of ""Patranomodon nyaphuli"".
"Patranomodon" have a short exposure of their palatine and premaxilla, which creates a shorter face compared to other anomodontons. This gives "Patranomodon" a shorter facial structure, shorter in length as well as small in size. They also have a reduced tabular, a slit-like interpterygoid vacuity, three sacral vertebrae, and a screw-shape jaw. The genus has many features indicating its herbivorous behavior: the division of the external adductor muscles in the jaw into two separate components, the medial and lateral side, as well as using a propalinal jaw movement while feeding on plant material.
Pentaceratops
The first specimens were collected by Charles Hazelius Sternberg in the San Juan Basin in New Mexico. Sternberg worked in commission for the Swedish Uppsala University. In 1921 he recovered a skull and a rump, specimens PMU R.200 and PMU R.286, at the Meyers Creek near the Kimbetoh Wash, in a layer of the Kirtland Formation, he sent the fossils to paleontologist Carl Wiman. In 1922 Sternberg decided to work on his own account and north of Tsaya Trading Post, in the Fossil Forest of San Juan County, he discovered a complete skeleton, which he sold to the American Museum of Natural History. The museum then sent out a team headed by Charles Mook and Peter Kaisen to assist Sternberg in securing this specimen; subsequent digging by Sternberg in 1923 brought the total of AMNH specimens to four. The rump of the main specimen was discarded by the museum because it had insufficient value as a display.
Evolution of dinosaurs
Sauropod fossils are found from the times of the earliest dinosaurs right up to the Cretaceous–Paleogene extinction event, from 227 to  Ma. Most sauropods are known from the Jurassic, to be more precise between 227 and 121 Ma.
Mylodon
Well preserved samples of "Mylodon" remains have been discovered in the Cueva del Milodón site in Patagonia, Chile along the southern flank of Cerro Benítez in the year 1896. Associated with bones of other early Patagonian animals, these remains of "Mylodon" date from an era earlier than 10,000 BC.
Deinonychus
The first remains were uncovered in 1931 in southern Montana near the town of Billings. The team leader, paleontologist Barnum Brown, was primarily concerned with excavating and preparing the remains of the ornithopod dinosaur "Tenontosaurus", but in his field report from the dig site to the American Museum of Natural History, he reported the discovery of a small carnivorous dinosaur close to a "Tenontosaurus" skeleton, "but encased in lime difficult to prepare." He informally called the animal "Daptosaurus agilis" and made preparations for describing it and having the skeleton, specimen AMNH 3015, put on display, but never finished this work. Brown brought back from the Cloverly Formation the skeleton of a smaller theropod with seemingly oversized teeth that he informally named "Megadontosaurus". John Ostrom, reviewing this material decades later, realized that the teeth came from "Deinonychus", but the skeleton came from a completely different animal. He named this skeleton "Microvenator".
Patagosaurus
In the 1970s many specimens of a previously unidentified dinosaur were found associated together in the same bed and locality: a pebbly stratum near a route to Cerro Condor. The specimens were first described by Jose Bonaparte in 1979. For the fossil he erected the genus "Patagosaurus", as well as its type species "P. fariasi". The generic name of "Patagosaurus" comes from the location of its find in Patagonia, and the fact that it is a reptile. The specific name honours Ricardo Farias, on whose land the initial discovery was made. The genus was originally known from an almost complete postcranial skeleton lacking a skull as the holotype, and many referred specimens however in 2003 it was found that a dentary was referable to the species, so more specimens are probably this taxon. Its skeleton was found near those of "Piatnitzkysaurus" and "Volkheimeria" in the Callovian to Oxfordian aged Patagonian deposits of the Cañadón Asfalto Formation. "Patagosaurus" is almost completely known with many articulated specimens found covering almost all of the skeleton, including parts of the skull. Over twelve specimens have been referred to the species, although some of the material is probably from a unique taxon. Bonaparte (1986) assigned three specimens other than the holotype PVL 4170, PVL 4076, MACN CH 934, and MACN CH 933 to the genus. While the holotype includes a post cranial skeleton, the others are known from cranial material and a nearly complete juvenile skeleton and skull. MACN CH 933 is directly comparable with the type material of "Patagosaurus", which confirms its association with the genus. A specimen first referred to "Patagosaurus" in 2003, MPEF-PV 1670 (which includes just a lower jaw), is also very similar to MACN CH 934, and differences can be associated with age, so therefore, MPEF-PV 1670 presumably represents adult cranial material. However, the teeth of MACN CH 934 are very different from those of both lower jaws (MACN CH 933 and MPEF-PV 1670), so it can be identified as another sauropod from the same deposit as "Patagosaurus". Thus, the taxon only certainly includes PVL 4170, MACN CH 933, and MPEF-PV 1670.
Aurorazhdarcho
A specimen originally classified as "Pterodactylus micronyx" (now "Aurorazhdarcho micronyx") was one of the earliest, if not the earliest, documented pterosaur fossil ever found. The holotype specimen of "P. micronyx", also known as the "Pester Exemplar", was originally part of the private fossil collection held by Archduchess Maria Anna of Austria. Evidence suggest that the Pester Exemplar was unearthed at some point between 1757, when Maria Anna was recovering from serious cases of pneumonia and tuberculosis, after which she began collecting fossils, and 1779, when the specimen was first studied by scientists. This overlaps with the possible time of discovery of the holotype specimen of "Pterodactylus antiquus", often considered the first pterosaur found, which was unearthed sometime between 1767 and 1784.
Georges Cuvier
In 1800 and working only from a drawing, Cuvier was the first to correctly identify in print, a fossil found in Bavaria as a small flying reptile, which he named the "Ptero-Dactyle" in 1809, (later Latinized as "Pterodactylus antiquus")—the first known member of the diverse order of pterosaurs. In 1808 Cuvier identified a fossil found in Maastricht as a giant marine lizard, the first known mosasaur.
Tenontosaurus
The first "Tenontosaurus" fossil was found in Big Horn County, Montana by an American Museum of Natural History (AMNH) expedition in 1903. Subsequent digs in the same area during the 1930s unearthed 18 more specimens, and four specimens were found during the 1940s. Despite the large number of fossil specimens, the animal was not named or scientifically described during this time, though Barnum Brown of the AMNH gave it the informal name "Tenantosaurus", "sinew lizard", in reference to the extensive system of stiffening tendons in its back and tail.
Megalosaurus
The presumption that carnivorous dinosaurs, like "Megalosaurus", were quadrupeds was first challenged by the find of "Compsognathus" in 1859. That, however, was a very small animal, the significance of which for gigantic forms could be denied. In 1870, near Oxford, the type specimen of "Eustreptospondylus" was discovered - the first reasonably intact skeleton of a large theropod. It was clearly bipedal. Shortly afterwards, John Phillips created the first public display of a theropod skeleton in Oxford, arranging the known "Megalosaurus" bones, held by recesses in cardboard sheets, in a more or less natural position. During the 1870s, North American discoveries of large theropods, like "Allosaurus", confirmed that they were bipedal. The Oxford University Museum of Natural History display contains most of the specimens from the original description by Buckland.
Strathcona Fiord
The first High Arctic terrestrial fossil vertebrates were discovered in 1975, in the Strathcona Fiord by a team led by Mary Dawson from the Carnegie Museum. These earliest finds include the fossil remains of an alligator, considered to be "Allognathosuchus", and also small arboreal mammals called "plagiomenids". Since then, field expeditions in Strathcona Fiord have yielded a much more complete picture of the biodiversity of the Eocene Arctic. Although, Eocene fossil vertebrates are known from other areas on Ellesmere Island (e.g., Stenkul Fiord), and Axel Heiberg Island, Strathcona Fiord has yielded the richest vertebrate fossil record.
Spinosauridae
The first spinosaurid fossil, a single conical tooth, was discovered circa 1820 by British paleontologist Gideon Mantell in the Wadhurst Clay Formation. In 1841 Sir Richard Owen mistakenly assigned it to a crocodilian he named "Suchosaurus" ("crocodile lizard"), a second species, "S. girardi", was later named in 1897. However, the spinosaurid nature of "Suchosaurus" was not recognized until a 1998 redescription of "Baryonyx."
Temnodontosaurus
The species "Temnodontosaurus platyodon" was named by William Conybeare in 1822 from the specimen BMNH 2003 from the Lyme Regis. This specimen is located at the British Museum of Natural History. "T. platyodon" is from the Upper Hettangian-Lower Sinemurian. It is the type species for "Temnodontosaurus"." T. platyodon" specimens have been found in England, Germany and Belgium. This includes the Lyme Regis in Dorset England, Herlikofen in Germany and Arlon in Belgium. Only one complete skeleton of "T. platyodon" is known of (BMNH 2003) and there is also a well preserved skull (BMNH R1158).
In 1995, Christopher McGowan explained that the previously named "T. risor" specimens are really juvenile versions of "T. platyodon". The specimen he used to back up his argument was one collected by David Sole in 1987 from Black Ven (East of Lyme Regis). The previously known "T. risor" specimens (three skulls) were seen previously as different to the "T. platyodon" species because they had larger orbits, smaller maxillae and curved snouts. However, McGowan described them as juveniles because of the small size of the forefin relative to the skull. The "T. risor" skulls are thought to be juveniles because the skull is relatively long compared to the postcranial skeleton.
Streptospondylus
In 1808, Cuvier scientifically described the theropod vertebrae as the first dinosaur remains ever. However, he considered them to be crocodilian and associated them with fossils of the Teleosauridae and the Metriorhynchidae. In 1822, Cuvier by the work of Henry De La Bèche became aware that these finds were very disparate, stemming from different periods. He abstained from naming them but in 1824 concluded that there were two main types. In 1825 Étienne Geoffroy Saint-Hilaire accordingly named two crocodilian skulls as the genus "Steneosaurus", the one, specimen MNHN 8900, becoming "Steneosaurus rostromajor", the other, MNHN 8902, "S. rostrominor".
Philippe-Charles Schmerling
In 1829 he discovered the first Neanderthal fossil in a cave in Engis, the partial cranium of a small child, although it was not recognized as such until 1936, and is now thought to be between 30,000-70,000 years old. It was the second discovery of a fossil of the genus Homo after the discovery of the Red Lady of Paviland in Wales in 1823.
Aurorazhdarcho
In 1856, Hermann von Meyer illustrated the Pester Exemplar, and designated it the type specimen of his new species "Pterodactylus micronyx" after comparison with the newly recognized and better-preserved pterosaur specimens, all of which were lumped together at that time into the single genus "Pterodactylus". However, by 1960, Peter Wellnhofer was not able to locate the specimen in the Hungarian National Museum or in the collections of Eötvös Loránd University. Wellnhofer therefore considered the specimen lost, and designated a neotype. The Pester Exemplar was re-discovered in 1982, during a re-organization of the collections in Eötvös Loránd University, and so the status of type specimen for "P. micronyx" reverted to it.
Pterosaur Beach
Pterosaur Beach is the name that paleontologists have given to an area along a long gone lagoon in what is now southwestern France. The area is notable because it is the first time that the fossil footprints of a landing pterosaur have been discovered. The fossil footprints are approximately 140 million years old.
Dimetrodon
Fossils now attributed to "Dimetrodon" were first studied by American paleontologist Edward Drinker Cope in the 1870s. Cope had obtained the fossils along with those of many other Permian tetrapods (four-limbed vertebrates) from several collectors who had been exploring a group of rocks in Texas called the Red Beds. Among these collectors were Swiss naturalist Jacob Boll, Texas geologist W. F. Cummins, and amateur paleontologist Charles Hazelius Sternberg. Most of Cope's specimens went to the American Museum of Natural History or to the University of Chicago's Walker Museum (most of the Walker fossil collection is now housed in the Field Museum of Natural History).
Pteranodon
Adult male "Pteranodon" were among the largest pterosaurs, and were the largest flying animals known until the late 20th century, when the giant azhdarchid pterosaurs were discovered. The wingspan of an average adult male "Pteranodon" was . Adult females were much smaller, averaging in wingspan. The largest specimen of "Pteranodon longiceps" from the Niobrara Formation measured from wingtip to wingtip. An even larger specimen is known from the Pierre Shale Formation, with a wingspan of , though this specimen may belong to the distinct genus and species "Geosternbergia maysei". While most specimens are found crushed, enough fossils exist to put together a detailed description of the animal.
Pteranodon
The diet of "Pteranodon" is known to have included fish; fossilized fish bones have been found in the stomach area of one "Pteranodon", and a fossilized fish bolus has been found between the jaws of another "Pteranodon", specimen AMNH 5098. Numerous other specimens also preserve fragments of fish scales and vertebrae near the torso, indicating that fish made up a majority of the diet of "Pteranodon" (though they may also have taken invertebrates).
Bone marrow
The earliest fossilised evidence of bone marrow was discovered in 2014 in "Eusthenopteron", a lobe-finned fish which lived during the Devonian period approximately 370 million years ago. Scientists from Uppsala University and the European Synchrotron Radiation Facility used X-ray synchrotron microtomography to study the fossilised interior of the skeleton's humerus, finding organised tubular structures akin to modern vertebrate bone marrow. "Eusthenopteron" is closely related to the early tetrapods, which ultimately evolved into the land-dwelling mammals and lizards of the present day.
Pteranodon
Other distinguishing characteristics that set "Pteranodon" apart from other pterosaurs include narrow neural spines on the vertebrae, plate-like bony ligaments strengthening the vertebrae above the hip, and a relatively short tail in which the last few vertebrae are fused into a long rod. The entire length of the tail was about 3.5% as long as the wingspan, or up to in the largest males.
Stenonychosaurus
A more complete skeleton of "Stenonychosaurus" was described by Dale Russell in 1969 from the Dinosaur Park Formation, which eventually formed the scientific foundation for a famous life-sized sculpture of "Stenonychosaurus" accompanied by its fictional, humanoid descendant, the "dinosauroid". "Stenonychosaurus" became a well-known theropod in the 1980s, when the feet and braincase were described in more detail. Along with "Saurornithoides", it formed the family Saurornithoididae. Based on differences in tooth structure, and the extremely fragmentary nature of the original "Troodon formosus" specimens, saurornithoidids were thought to be close relatives while "Troodon" was considered a dubious possible relative of the family. Phil Currie, reviewing the pertinent specimens in 1987, showed that supposed differences in tooth and jaw structure among troodontids and saurornithoidids were based on age and position of the tooth in the jaw, rather than a difference in species. He reclassified "Stenonychosaurus inequalis" as well as "Polyodontosaurus grandis" and "Pectinodon bakkeri" as junior synonyms of "Troodon formosus". Currie also made Saurornithoididae a junior synonym of Troodontidae. In 1988, Gregory S. Paul went farther and included "Saurornithoides mongoliensis" in the genus "Troodon" as "T. mongoliensis", but this reclassification, along with many other unilateral synonymizations of well known genera, was not adopted by other researchers. Currie's classification of all North American troodontid material in the single species "Troodon formosus" became widely adopted by other paleontologists, and all of the specimens once called "Stenonychosaurus" were referred to as "Troodon" in the scientific literature through the early 21st century.
Besanosaurus
The bones of "Besanosaurus" were first discovered in "Sasso Caldo" quarry in the spring of 1993 by the volunteers of the paleontological group of Besano, a small town in the Lombardy region of north Italy. The fossil was almost completely embedded in the rock and could be first seen only through x-rays; to detect the content of the 38 slabs of stone enclosing the skeleton, 145 radiographs were necessary. The skeleton of "Besanosaurus" came to light in the paleontological laboratory of Museo Civico di Storia Naturale di Milano after 16500 hours of preparation. The preparators removed the rock enclosing the fossil step by step, working under a stereo microscope with chisels, needles and pins. The 38 slabs were reassembled and a silicon rubber matrix was made, from which a cast of the original specimen was obtained.
Pterodactyloidea
In 2014, fossils from the Shishugou Formation of China were classified as the most basal pterodactyloid yet found, "Kryptodrakon". At a minimum age of about 161 my, it is about 5 million years older than the oldest previously known confirmed specimens. Previously, a fossil jaw recovered from the Middle Jurassic Stonesfield Slate formation in the United Kingdom, was considered the oldest known. This specimen supposedly represented a member of the family Ctenochasmatidae, though further examination suggested it belonged to a teleosaurid stem-crocodilian instead of a pterosaur. O'Sullivan and Martill (2018) described a partial synsacrum from the Stonesfield Slate identified as possibly pterodactyloid based on the number of incorporated sacrals although they commented that the morphology was perhaps closer to that of wukongopterids. If correctly identified, it would be the oldest pterodactyloid fossil known.
Polyonax
During an 1873 trip through the western US, paleontologist and naturalist Edward Drinker Cope collected some fragmentary dinosaurian material which he soon named as a new genus. Catalogued today as AMNH FR 3950, the type material included three dorsal vertebrae, limb bone material, and what are now known to be horn cores, from a subadult individual. Although it was briefly mixed up with hadrosaurs, and even considered to be a possible synonym of "Trachodon", it was recognized as a horned dinosaur in time for the first monograph on horned dinosaurs (1907), wherein it was regarded as based on indeterminate material. Today, the name is used as little more than a historical curiosity, as it dates from a time before horned dinosaurs were known to exist. The most recent review listed it as an indeterminate ceratopsid.
Timeline of pterosaur research
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.
Pisanosaurus
"Pisanosaurus" is known from a single fragmented skeleton discovered in 1962 by Galileo Juan Scaglia at the Agua de Las Catas locality in the Ischigualasto Formation, in La Rioja, Argentina. It is based on a specimen given the designation PVL 2577, which consists of a partial skull with a fragmentary right maxilla with teeth, and incomplete right mandibular ramus (lower jaw), six incomplete cervical vertebrae, seven incomplete dorsal vertebrae, molds of five sacral vertebrae, a rib and several rib fragments, a fragmentary right scapula, a coracoid, molds of a fragmentary ilium, ischium and pubic bone, an impression of three metacarpals, the complete femora, the right tibia, the right fibula, with an articulated astragalus and calcaneum, a tarsal element with a metatarsal, metatarsals III and IV, three phalanges from the third toe and five phalanges, among them the ungual, from the fourth toe, and an indeterminate long bone fragment.
Pteranodontia
Pteranodontia was originally named by O.C. Marsh in 1876. In 2003, it was given a phylogenetic definition by David Unwin as the common ancestor of "Pteranodon" and "Nyctosaurus" plus all its descendants. Though Marsh had originally named this group based on the shared absence of teeth in those species, most analyses show that all of the traditional "ornithocheiroid" pterosaurs are also members of this clade.
Cetiosauriscus
The sauropod fossil today known as "Cetiosauriscus stewarti" was discovered in May 1898 by clay workers in the area around Fletton to the south of Peterborough and east of the Great Northern Railway. Pits in this region expose the fossil-rich sedimentary rocks of the marine Oxford Clay, which is of middle Callovian age and today regarded as one of the classic geological formations of British palaeontology. The sauropod fossil possibly stems from NPBCL pit No. 1, which was the northernmost pit operated by the New Peterborough Brick Company Limited, and which produced the most vertebrate fossils. The discovery was brought to the attention of Leeds, who, after excavation, took the sauropod specimen to Eyebury, the Leeds' family home. In mid-August, after some cleaning and repairing of the specimen, geologist Henry Woodward visited Eyebury and produced a life-sized drawing of the remains for presentation at the British Association for the Advancement of Science Meeting. Following this presentation, on 17 August 1898, Henry Woodward returned with American palaeontologist Othniel Charles Marsh, who considered the sauropod to be closely related to the North American taxon "Diplodocus". Alfred Leeds offered the sauropod to the British Museum of Natural History (BMNH, now abbreviated as NHMUK) for £250, which would equate to about £30,529 in 2017. The NHMUK had earlier in 1890 and 1892 bought the First and Second Collections of Alfred Leeds, respectively. Woodward, Keeper of Geology at the NHMUK, had "great pleasure" to recommend to the Trustees of the NHMUK the fossil be purchased. The purchase was sanctioned on 25 February 1899, along with the purchase of assorted other remains for just over £357 (~£43,596 now), where the Leeds sauropod gained the accession number BMNH R3078 (now NHMUK R3078).
Panthera leo fossilis
Panthera leo fossilis is a fossil cat of the genus "Panthera", which was first excavated near Mauer in Germany, and lived during the Upper Pleistocene.
Bone fragments of "P. l. fossilis" were also excavated near Pakefield in the United Kingdom, which are estimated at 680,000 years old.
Bone fragments excavated near Isernia in Italy are estimated at between 600,000 and 620,000 years old.
The first Asian record of a "fossilis" lion was found in the Kuznetsk Basin in western Siberia and dates to the late Early Pleistocene.
Stenonychosaurus
In 1982, Dale A. Russell, then curator of vertebrate fossils at the National Museum of Canada in Ottawa, conjectured a possible evolutionary path for "Stenonychosaurus", if it had not perished in the Cretaceous–Paleogene extinction event 65 million years ago, suggesting that it could have evolved into intelligent beings similar in body plan to humans. Over geologic time, Russell noted that there had been a steady increase in the encephalization quotient or EQ (the relative brain weight when compared to other species with the same body weight) among the dinosaurs. Russell had discovered the first Troodontid skull, and noted that, while its EQ was low compared to humans, it was six times higher than that of other dinosaurs. Russell suggested that if the trend in "Stenonychosaurus" evolution had continued to the present, its brain case could by now measure 1,100 cm, comparable to that of a human.
Geosternbergia
The first "G. sternbergi" fossil was collected by George F. Sternberg in 1952 and described by John Christian Harksen in 1966, from the lower portion of the Niobrara Formation. It was older than "Pteranodon longiceps" and is considered by Bennett to be the direct ancestor of that species.
Platypus
The oldest discovered fossil of the modern platypus dates back to about 100,000 years ago, during the Quaternary period. The extinct monotremes "Teinolophos" and "Steropodon" were once thought to be closely related to the modern platypus, but are now considered more basal taxa. The fossilised "Steropodon" was discovered in New South Wales and is composed of an opalised lower jawbone with three molar teeth (whereas the adult contemporary platypus is toothless). The molar teeth were initially thought to be tribosphenic, which would have supported a variation of Gregory's theory, but later research has suggested, while they have three cusps, they evolved under a separate process. The fossil is thought to be about 110 million years old, making it the oldest mammal fossil found in Australia. Unlike the modern platypus (and echidnas), "Teinolophos" lacked a beak.
Parrot
The first uncontroversial parrot fossils date to tropical Eocene Europe around 50 mya. Initially, a neoavian named "Mopsitta tanta", uncovered in Denmark's Early Eocene Fur Formation and dated to 54 mya, was assigned to the Psittaciformes. However, the rather nondescript bone is not unequivocally psittaciform, and it may rather belong to the ibis genus "Rhynchaeites", whose fossil legs were found in the same deposits.
Several fairly complete skeletons of parrot-like birds have been found in England and Germany. These are probably not transitional fossils between ancestral and modern parrots, but rather lineages that evolved parallel to true parrots and cockatoos:
The earliest records of modern parrots date to around 23–20 mya. The fossil record—mainly from Europe—consists of bones clearly recognisable as belonging to anatomically modern parrots. The Southern Hemisphere contains no known parrot-like remains earlier than the Early Miocene around 20 mya.
Elizabeth Philpot
The Philpot sisters' extensive and meticulously labelled fossil collection was used for research by many geologists. All three sisters contributed to the collection, but it was Elizabeth Philpot who corresponded with leading geologists like William Buckland, William Conybeare, and Henry De la Beche about the collection. The collection was well known for its fossil fish. It also contained the fossil teeth that William Buckland combined with a famous partial skeleton discovered by Mary Anning when he described the pterosaur "Pterodactylus macronyx" (later renamed by Richard Owen to "Dimorphodon macronyx"), in 1829. In his famous 1824 paper that described the almost complete plesiosaur skeleton discovered by Anning in 1823 Conybeare mentioned examining a plesiosaur skull in the possession of "Miss Philpot".
Serradraco
In 1874, Richard Owen named a pair of lower jaws from the collection of Samuel Husband Beckles, found at St Leonards-on-Sea in Sussex, as a new species of "Pterodactylus": "Pterodactylus sagittirostris". The specific name means "arrowhead-snouted" in Latin, referring to the mandible profile in upper view. In 1888, Edwin Tulley Newton, conforming to the soon to be published pterosaur systematics by Richard Lydekker, renamed the species into "Ornithocheirus sagittirostris". In July 1891, the British Museum (Natural History), the present Natural History Museum, bought the piece from the heirs of Beckles.
Pteranodon
In 2010, pterosaur researcher Alexander Kellner revisited H.W. Miller's classification. Kellner followed Miller's opinion that the differences between the "Pteranodon" species were great enough to place them into different genera. He placed "P. sternbergi" into the genus named by Miller, "Geosternbergia", along with the Pierre Shale skull specimen which Bennett had previously considered to be a large male "P. longiceps". Kellner argued that this specimen's crest, though incompletely preserved, was most similar to "Geosternbergia". Because the specimen was millions of years younger than any known "Geosternbergia", he assigned it to the new species "Geosternbergia maysei". Numerous other pteranodont specimens are known from the same formation and time period, and Kellner suggested they may belong to the same species as "G. maysei", but because they lack skulls, he could not confidently identify them.
History of paleontology in the United States
The year after the "Hadrosaurus"'s fossils were first identified, 1859, state agricultural chemist Philip T. Tyson found the first documented dinosaur fossils of the Arundel Formation in an iron pit at Bladensburg, Maryland. The discovery consisted of two fossil teeth. Tyson took the dinosaur teeth to a local doctor named Christopher Johnston. Johnston cut thin sections of one tooth to examine it under a microscope. Johnson named the teeth "Astrodon". In 1865 Joseph Leidy formally named the species "Astrodon johnstoni" after Christopher Johnston. This represents the first formal naming of a sauropod species in North America.
Paleontology in Pennsylvania
One of the earliest notable events in Pennsylvania paleontology was the October 5th, 1787 presentation by Caspar Wistar and Timothy Matlack of a probable dinosaur metatarsal discovered in Late Cretaceous rocks near Woodbury Creek in New Jersey as "'a large thigh bone'" to the American Philosophical Society in Philadelphia. During the industrial revolution, Carboniferous-aged coal deposits in Pennsylvania were the sites of serendipitous discoveries of early fossil tetrapod trackways. Such discoveries generally occur when the excavation of coal mines removes the rock underlying the trackway, leaving it exposed on the tunnel's ceiling. Later, during the 1840s, Charles Lyell examined some local purported fossil bird and mammal tracks and found that they were actually petroglyphs left by local indigenous people. Depictions of trace fossils in indigenous rock art are known from throughout North America.
Pterodactylus
It was the German/French scientist Johann Hermann who first stated that "Pterodactylus" used its long fourth finger to support a wing membrane. In March 1800, Hermann alerted the French scientist George Cuvier to the existence of Collini's fossil, believing that it had been captured by the occupying armies of Napoleon and sent to the French collections in Paris (and perhaps to Cuvier himself) as war booty; at the time special French political commissars systematically seized art treasures and objects of scientific interest. Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined it personally), which he believed to be a mammal, including the first known life restoration of a pterosaur. Hermann restored the animal with wing membranes extending from the long fourth finger to the ankle and a covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added a membrane between the neck and wrist, as is the condition in bats. Cuvier agreed with this interpretation, and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very short description. Cuvier remarked, "[It is not possible to doubt that the long finger served to support a membrane that, by lengthening the anterior extremity of this animal, formed a good wing.]" However, contrary to Hermann, Cuvier was convinced the animal was a reptile.
Pectinodon
Pectinodon is a genus of dinosaurs from the Late Cretaceous period (66 mya). It currently contains a single valid species, Pectinodon bakkeri (sometimes classified as "Troodon bakkeri"), known mainly from teeth, as well as fragments of juvenile skeletons and eggshells.
Pteranodon
Historically, the terrestrial locomotion of "Pteranodon", especially whether it was bipedal or quadrupedal, has been the subject of debate. Today, most pterosaur researchers agree that pterosaurs were quadrupedal, thanks largely to the discovery of pterosaur trackways.
Eudimorphodon
Eudimorphodon was a pterosaur that was discovered in 1973 by Mario Pandolfi in the town of Cene, Italy and described the same year by Rocco Zambelli. The nearly complete skeleton was retrieved from shale deposited during the Late Triassic (mid to late Norian stage), making "Eudimorphodon" one of the oldest pterosaurs known. It had a wingspan of about and at the end of its long bony tail may have been a diamond-shaped flap like in the later "Rhamphorhynchus". If so, the flap may have helped it steer while maneuvering in the air. "Eudimorphodon" is known from several skeletons, including juvenile specimens.
Rhamphinion
The type species, "Rhamphinion jenkinsi", was described and named in 1984 by Kevin Padian, based on holotype MNA V 4500, a partial skull including the occipital region, a partial left jugal, a fragment of the lower jaw including two preserved teeth and the impression of a third, and one other fragment that could not be identified. The specific name honors the discoverer of the fossil, Farish Alston Jenkins Jr. At the time, it was the oldest known pterosaur specimen from the Western Hemisphere (now overtaken by "Caelestiventus"). Padian did not assign it to any family or suborder within Pterosauria, but did note that the jugal was unlike that of pterodactyloids, and so may have belonged to a "rhamphorhynchoid", i.e. a basal pterosaur. A bone from the wing of another pterosaur from the same formation, found in 1981, may have also belonged to a "rhamphorhynchoid", with a wingspan of about 1.5 m (4.9 ft). Peter Wellnhofer agreed that a "rhamphorhynchoid" identity was very likely, but David Unwin was more hesitant to classify its fragmentary remains in "The Pterosaurs: From Deep Time," noting it only as a "possibly valid species of uncertain relationships." A recent cladistic analysis of "Jidapterus" recovers "Rhamphinion" as a close relative of "Dimorphodon" and "Parapsicephalus".
Peramelemorphia
Many specimens of modern peramelemorphian (e.g. "Perameles" spp. and "Isoodon" spp.) have been recovered in the fossil record from Pleistocene and Holocene fossil localities. However, very few fossil species have been recovered to date. The first species of fossil peramelemorphian was described by R. A. Stirton in 1955. The specimen Stirton described was a partial lower jaw from the Tirari Desert in Central Australia, Pliocene in age. The lower jaw morphology suggested a relationship with bilbies (Family Thylacomyidae), and was named "Ischnodon australis".
Nothronychus
The first fossil evidence later attributed to "Nothronychus" was discovered by a team of paleontologists working in the Zuni Basin of New Mexico at the Haystack Butte site. A therizinosaur ischium (a hip bone) had originally been mistaken for a squamosal, a part of the skull crest of the newly discovered ceratopsian "Zuniceratops". However, closer examination revealed the true identity of the bone, and soon more parts of the skeleton were found. The New Mexico team, led by paleontologists Jim Kirkland and Doug Wolfe, published their find in the "Journal of Vertebrate Paleontology" on 22 August 2001, making it the type specimen of the new species "Nothronychus mckinleyi". The "Arizona Republic" newspaper, however, was first to announce the name on 19 June 2001, in a column by R.E. Molnar. The generic name is derived from Greek νωθρός, "nothros", "heavy" or "slothful", and ὄνυξ, "onyx", "claw". The specific name honours rancher Bobby McKinley on whose land the find was made. The holotype, specimen MSM P2106, consists of two skull fragments, a braincase, some vertebrae and parts of the shoulder girdle, forelimbs, pelvis and hindlimbs.
A second, more complete therizinosaur specimen, UMNH VP 16420, was discovered from the Tropic Shale formation (dating to the early Turonian stage) of southern Utah in 2000 by Merle Graffam, a resident of Big Water, Utah. The area around Big Water had been subject to several expeditions by teams from the Museum of Northern Arizona (MNA), and was known for its abundance of marine reptile fossils, especially plesiosaurs. During part of the late Cretaceous period, the region had been submerged under a shallow sea, the Western Interior Seaway, and preserves extensive marine deposits. Graffam's initial discovery (a large, isolated toe bone) came as a surprise to scientists, as it clearly belonged to a land-dwelling dinosaur, rather than a plesiosaur. However, the location of the bone at the time would have been nearly 100 kilometers from the Cretaceous shoreline. An excavation of the area by an MNA crew revealed more of the skeleton, and the scientists found that it was a therizinosaur, and the first example of that group to be found in the Americas. All previous therizinosaur fossils had come from China and Mongolia. It is the most complete therizinosaurid specimen known but lacks the skull.
Stenonychosaurus
Dinosaur eggs and nests were discovered by John R. Horner in 1983 in the Two Medicine Formation of Montana. Varriccho et al. (2002) have described eight of these nests found to date. These are all in the collection of the Museum of the Rockies and their accession numbers are MOR 246, 299, 393, 675, 676, 750, 963, 1139. Horner (1984) found isolated bones and partial skeletons of the hypsilophodont "Orodromeus" very near the nests in the same horizon and described the eggs as those of "Orodromeus". Horner and Weishampel (1996) reexamined the embryos preserved in the eggs and determined that they were those of "Troodon", not "Orodromeus". Varricchio et al. (1997) made this determination with even more certainty when they described a partial skeleton of an adult "Troodon" (MOR 748) in contact with a clutch of at least five eggs (MOR 750), probably in a brooding position. Van der Reest and Currie considered it possible that the Two Medicine troodont was the same species as "Stenonychosaurus".
Pterygotioidea
The first pterygotioid fossils to be uncovered were those of the type genus, "Pterygotus". Louis Agassiz, a Swiss-American biologist and geologist, described the fossils in 1839 and named it "Pterygotus" in the mistaken belief that they were the fossil remains of a large fish; ""Pterygotus"" translating to "winged fish". Agassiz first recognized the true nature of the fossils as arthropod remains five years later in 1844 after having examined more complete fossils recovered in the Old Red Sandstone of Scotland.
Paleontology in Utah
One of the earliest expeditions into Utah occurred during the summer and fall of 1870, when an expedition led by O. C. Marsh collected fossils in on behalf of Yale University. Later, in 1908, paleontologist Earl Douglass was excavating fossils on behalf of the Carnegie Museum of Natural History. Director Holland of the museum visited Douglass's camp that year. In September, Holland suggested that Douglass search for Jurassic dinosaur fossils in the Uinta Mountains north of his camp. Previous surveyors and paleontologists had noted dinosaur fossils of that age there and Holland suspected that they were promising hunting grounds. Douglass agreed and they set off to the Uinta Mountains the next day. Together they found a "Diplodocus" femur southeast of Split Mountain near the Green River. Encouraged by the discovery of the femur, Douglass returned to the Uinta Mountains the next year. Douglass had little luck during the spring and early part of the summer, but on August 17 he found a series of eight fossil "Apatosaurus" tail vertebrae, still articulated. On receiving word of the discovery, Holland returned to the site.
Pteranodon
"Pteranodon" species are extremely well represented in the fossil record, allowing for detailed descriptions of their anatomy and analysis of their life history. Over 1,000 specimens have been identified, though less than half are complete enough to give researchers good anatomical information. Still, this is more fossil material than is known for any other pterosaur, and it includes both male and female specimens of various age groups and possibly species.
Pteranodontoidea
Pteranodontoidea is an extinct clade of ornithocheiroid pterosaurs from the Early Cretaceous to the Late Cretaceous (middle Barremian to middle Campanian stages) of Asia, 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. Ornithocheiroidea is sometimes considered to be the senior synonym of Pteranodontoidea, however its depends on it definition. Ornithocheiroidea was originally defined as an apomorphy-based taxon by Christopher Bennett in 1994. Later, Kellner (2003) redefined it to represent the node of "Anhanguera", "Pteranodon", "Quetzalcoatlus" and "Dsungaripterus". Later, David Unwin (2003) suggested a different definition, the node that contains "Pteranodon longiceps" and "Istiodactylus latidens", thus making "Pteranodontoidea" a junior synonym of Ornithocheiroidea. Brian Andres (2008, 2010, 2014) in his analyses, converts Ornithocheiroidea using the definition of Kellner (2003) to avoid this synonymy.
Pterygotus
The first fossils of "Pterygotus" were found in deposits of Lochkovian-Pragian (Early Devonian) age by quarrymen in Scotland and western England, who referred to the large fossil remains as "Seraphims". Louis Agassiz, a Swiss-American biologist and geologist, described the fossils in 1839 and named it "Pterygotus" in the belief that they were the fossil remains of a large fish; ""Pterygotus"" translating to "winged fish". The specimens described by Agassiz from England were referred to a species he dubbed "Pterygotus problematicus". Agassiz first recognized the true nature of the fossils as arthropod remains five years later in 1844 after having examined more complete fossils recovered in the Old Red Sandstone of Scotland. Although recognizing the fossils of "Pterygotus" as arthropod was closer to its modern phylogenetical position, Agassiz would consider "Pterygotus" to represent a crustacean of the Entomostraca subclass. Although Frederick M'Coy did note that "Pterygotus" resembled the Limulidae and the previously discovered eurypterid "Eurypterus" in 1849, he classified both "Eurypterus" and "Pterygotus" as crustaceans.
Paleontology in the United States
By the beginning of the 19th century, their fossil footprints definitely had been found in Massachusetts and later, Connecticut. These tracks were researched by the Reverend Edward Hitchcock and instrumental to the establishment of ichnology, the study of trace fossils, as a science. During the late 1850s, the world's first reasonably complete dinosaur skeleton was discovered in New Jersey. Joseph Leidy would name it Hadrosaurus. This was the first known dinosaur, the first dinosaur to be interpreted as two-legged, and the first to be mounted for exhibition in a museum. Later in the century Edward Drinker Cope described a new carnivorous dinosaur from New Jersey he called Laelaps. When he gave fellow paleontologist Othniel Charles Marsh a tour of the quarry the fossils were taken from Marsh secretly convinced Cope's workers to send fossils to himself instead. Marsh also learned that the name Laelaps had already been used for a different animal and renamed Cope's dinosaur Dryptosaurus out from under him. This instigated a bitter rivalry to collect the most fossils and name the most new prehistoric species that took the two paleontologists and their crews deep into the west looking for fossils and attempting to undermine each other's fieldwork.
Cuvieronius
The species now known as "Cuvieronius hyodon" was among the first fossil animals from the New World to be studied. The first remains of this species were recovered from Ecuador by Alexander von Humboldt, at a location the local population referred to as the "Field of Giants". Humboldt recognized that, rather than being bones of giant humans as had been thought by the local population and previous Spanish colonists, they were similar to the giant elephants ("Mastodon") being described from Ohio. Humboldt sent teeth that he had collected from Mexico, Ecuador, and Chile to French anatomist Georges Cuvier, who classified the teeth into two species, which he referred to as the "mastodonte des cordilières" and the "mastodonte humboldtien", in an 1806 paper. It was not until 1824 that Cuvier formally named the species. He referred both to the genus "Mastodon", calling them "M. andium" and "M. humboldtii".
Stonesfield
The mines were also one of Britain's richest sources of Middle Jurassic vertebrate fossils. The first fossil bones to be identified as those of a dinosaur were found early in the 19th century near Stonesfield. They are part of the skeleton of a bipedal carnivore, and in 1824 the pioneering palaeontologist William Buckland named it "Megalosaurus". The bones are now displayed in the Oxford University Museum of Natural History. Other reptiles found at Stonesfield include the crocodile "Steneosaurus", pterosaur "Rhamphocephalus" and the type specimen of the theropod genus "Iliosuchus".
Istiodactylus
Istiodactylus is a genus of pterosaur that lived during the Early Cretaceous period, about 120 million years ago. The first fossil was discovered on the Isle of Wight in England, and became the holotype specimen of a new species, "O. latidens" (Latin for "wide tooth"), in the genus "Ornithodesmus" in 1901. This species was moved to its own genus, "Istiodactylus", in 2001; this name is Greek for "sail finger". More specimens were described in 1913, and "Istiodactylus" was the only pterosaur known from three-dimensionally preserved fossils for much of the 20th century. In 2006, a species from China, "I. sinensis", was assigned to "Istiodactylus", but it has since been found to belong to a different genus.
Timeline of pterosaur research
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. 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. The British anatomist Sir Richard Owen dubbed this vanished order the Pterosauria. Soon after, he described Britain's own first pterosaur, "Dimorphodon". 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.
Paleontology in Kansas
Paleontology in Kansas refers to paleontological research occurring within or conducted by people from the U.S. state of Kansas. Kansas has been the source of some of the most spectacular fossil discoveries in US history. The fossil record of Kansas spans from the Cambrian to the Pleistocene. From the Cambrian to the Devonian, Kansas was covered by a shallow sea. During the ensuing Carboniferous the local sea level began to rise and fall. When sea levels were low the state was home to richly vegetated deltaic swamps where early amphibians and reptiles lived. Seas expanded across most of the state again during the Permian, but on land the state was home to thousands of different insect species. The popular pterosaur "Pteranodon" is best known from this state. During the early part of the Cenozoic era Kansas became a savannah environment. Later, during the Ice Age, glaciers briefly entered the state, which was home to camels, mammoths, mastodons, and saber-teeth. Local fossils may have inspired Native Americans to regard some local hills as the homes of sacred spirit animals. Major scientific discoveries in Kansas included the pterosaur "Pteranodon" and a fossil of the fish "Xiphactinus" that died in the act of swallowing another fish.
Astrodon
Two dinosaur teeth were received in late November 1858 by chemist Philip Thomas Tyson from John D. Latchford. They had been found in Latchford's open iron ore pit in the Arundel Formation at Swampoodle near Muirkirk in Prince George's County, Maryland. Tyson let them be studied by the dentist Christopher Johnston, professor at the Baltimore Dental College, who cut one tooth in half and thereby discovered a characteristic star-formed cross-section. Johnston named "Astrodon" in 1859. However, he did not attach a specific epithet, so Joseph Leidy is credited with naming "Astrodon johnstoni" (the type species) in 1865, with as holotype specimen YPM 798. If Johnston had attached a specific epithet, it would have been the second dinosaur species identified in the United States. Johnston incorrectly stated that the site of the discovery had been an iron mine near the town of Bladensburg.
Rodan (film)
Shigeru confirms that the creature he saw did indeed resemble a "Pteranodon", and that it had eaten all of the Meganulon. He and a group of police and scientists once again descend into the mine and enter the cave where the egg had been. They are able to recover a fragment of the shell before a rock slide forces them to flee back to the surface. In the lab, Dr. Kashiwagi is able to determine the size of the egg and its age: 200 million years old. After amassing the evidence, Kashiwagi calls a meeting with members of the town, along with members of the Japanese Self-Defence Force, to communicate his findings. He tells the men that the UFO seen flying all across the world at supersonic speeds is a gigantic pterosaur he has dubbed Rodan. How Rodan could have resurfaced after millions of years is also a mystery, but Kashiwagi theorizes that nuclear bomb testing, which loosened the Earth's crust and opened cavities to long buried crevices and caves, might be the possible cause.
Spinosauridae
The first fossils referred to a spinosaurid were discovered in 1912 at the Bahariya Formation in Egypt. Consisting of dorsal vertebrae, skull fragments, and teeth, these remains became the holotype specimen of the new genus and species "Spinosaurus aegyptiacus" in 1915, when they were described by German paleontologist Ernst Stromer. The dinosaur's name meant "egyptian spine lizard", in reference to the unusually long neural spines not seen previously in any other theropod. In April 1944, the holotype of "S. aegyptiacus" was destroyed during an allied bombing raid in World War II. In 1934, Stromer referred a partial skeleton also from the Bahariya Formation to a new species of "Spinosaurus", this specimen has since been alternatively assigned to another African spinosaurid, "Sigilmassasaurus."
Aerodactylus
The fossil remains of this species have been found only in the Solnhofen limestone of Bavaria, Germany, dated to the late Jurassic Period (early Tithonian), about 150.8–148.5 million years ago. Like all pterosaurs, the wings of "Aerodactylus" were formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by collagen fibres and externally by keratinous ridges. Several well preserved fossils have shown that "Aerodactylus" was covered in a short, dense coat of bristly pycnofibres, and that it had a rounded triangular crest on its head, as well as a backward-pointing lappet. It is named after the Pokémon Aerodactyl. The validity of "Aerodactylus" has been disputed, with some pterosaur experts suggesting that none of the specimens referred to this genus are distinguishable from "Pterodactylus".
Carnotaurus
"Carnotaurus" was the first theropod dinosaur discovered with a significant number of fossil skin impressions. These impressions, found beneath the skeleton's right side, come from different body parts, including the lower jaw, the front of the neck, the shoulder girdle, and the rib cage. The largest patch of skin corresponds to the anterior part of the tail. Originally, the right side of the skull also was covered with large patches of skin—this was not recognized when the skull was prepared, and these patches were accidentally destroyed. Still, the surface texture of much of the right side of the skull is very different from that of the left side, and probably shows some features of the scalation pattern of the head.
Phanerozoic
The Proterozoic-Phanerozoic boundary is at  million years ago. In the 19th century, the boundary was set at time of appearance of the first abundant animal (metazoan) fossils but several hundred groups (taxa) of metazoa of the earlier Proterozoic era have been identified since the systematic study of those forms started in the 1950s. Most geologists and paleontologists would probably set the Proterozoic-Phanerozoic boundary either at the classic point where the first trilobites and reef-building animals (archaeocyatha) such as corals and others appear; at the first appearance of a complex feeding burrow called "Treptichnus pedum"; or at the first appearance of a group of small, generally disarticulated, armored forms termed 'the small shelly fauna'. The three different dividing points are within a few million years of each other.
Matheronodon
In 2017, the fossil material was described by Godefroit along with Géraldine Garcia, Bernard Gomez, Koen Stein, Aude Cincotta, Ulysse Lefèvre, and Valentin in a research paper published by the journal "Nature Communications". They assigned this material to the genus "Matheronodon", with the name being formed from Philippe Matheron (who described the first dinosaur fossils found in Provence, or the southeastern region of France) and the suffix "-odon", a derivative of "odous" (Greek for "tooth"). They also named the type and only species "M. provincialis", with "provincialis" being Latin for Provence. For the study, a CT scan of the type maxilla was conducted at the University of Liège, and a thin section of an isolated maxillary tooth was examined under a microscope.
Thalassodromeus
The first known "Thalassodromeus" specimen was collected in 1983 near the town of Santana do Cariri in the Araripe Basin of northeastern Brazil. Found in outcrops of the Romualdo Formation, it was collected over a long period of time in several pieces. The specimen (catalogued as DGM 1476-R at the Museu de Ciências da Terra) was preserved in a calcareous nodule, and consists of an almost-complete, three-dimensional skull (pterosaur bones are often flattened compression fossils), missing two segments of the bottom of the skull and mandible and the front of the lower jaw. The left jugal region and right mandibular ramus (half of the mandible) are pushed slightly inward. The skull was first reported in a 1984 Italian book, and preliminarily described and figured in 1990 by palaeontologists Alexander W. A. Kellner and Diogenes de Almeida Campos. Although the pieces of skull had been divided between museums in South and North America, they were assembled before 2002.
In 2002, Kellner and Campos described and named the new genus and species "Thalassodromeus sethi", skull DGM 1476-R being the holotype specimen. The generic name is derived from the Ancient Greek words θάλασσα ("thálassa", "sea") and δρομεύς ("dromeús", "runner"), meaning "sea runner" in reference to the animal's supposed skim-feeding behaviour. The specific name refers to the ancient Egyptian god Seth. The specimen was not fully prepared at the time of this preliminary description. The original describers chose the name "sethi" because the crest of the pterosaur was supposedly reminiscent of the crown worn by Seth; however, the palaeontologists André Jacques Veldmeijer, Marco Signore, and Hanneke J. M. Meijer pointed out in 2005 that the crown (with its two tall plumes) was typically worn by the god Amon (or Amon-Ra) and his manifestations—not by Seth.
Weald
Many important fossils have been found in the sandstones and clays of the Weald, including, for example, "Baryonyx". The famous scientific hoax of Piltdown Man was claimed to have come from a gravel pit at Piltdown near Uckfield. The first "Iguanodon" was identified after a Mary Mantell unearthed some fossilised teeth by a road in Sussex in 1822. Her husband, Gideon Mantell, noticed they were similar to modern iguana teeth but many times larger; this important find led to the discovery of dinosaurs.
Pterodactylus
Pterodactylus ( , from the , ', meaning "winged finger") is an extinct genus of pterosaurs, whose members are popularly known as pterodactyls ( ). It is currently thought to contain only a single species, Pterodactylus antiquus, the first pterosaur species to be named and identified as a flying reptile.
Pteranodon
Methods used to estimate the mass of large male "Pteranodon" specimens (those with wingspans of about 7 meters) have been notoriously unreliable, producing a wide range of estimates from as low as to as high as . In a review of pterosaur size estimates published in 2010, researchers Mark Witton and Mike Habib demonstrated that the latter, largest estimates are almost certainly incorrect given the total volume of a "Pteranodon" body, and could only be correct if the animal "was principally aluminium". Witton and Habib considered the methods used by researchers who obtained smaller mass estimates equally flawed. Most have been produced by scaling modern animals such as bats and birds up to "Pteranodon" size, despite the fact that pterosaurs have vastly different body proportions and soft tissue anatomy from any living animal.
Aublysodon
The first skeletal material referred to an original "Aublysodon" species was a partial skull unearthed in Jordan, Montana in 1966 and described by Ralph Molnar in 1977/1978. The skull, specimen LACM 28741 at forty-five centimetres the length of an average human arm, bore pointed teeth attached to a long narrow snout. First thought to be a juvenile "Tyrannosaurus", then interpreted as a large dromaeosaurid, this "Jordan theropod" was given the name "Aublysodon molnaris" by Gregory S. Paul in 1988; in 1990 the name was by Paul emended to "Aublysodon molnari", applying the correct genitive. It was made a separate genus "Stygivenator" by George Olshevsky in 1995, but was later, in 2004, reinterpreted to be a juvenile "Tyrannosaurus rex" by Thomas Carr and Tom Williamson. Another partial skeleton from New Mexico, specimen OMNH 10131, was in 1990 considered to represent "Aublysodon", but later research by Thomas Carr and Tom Williamson first referred it to "Daspletosaurus" and ultimately to "Bistahieversor". In 1988 Paul also created another species when renaming "Shanshanosaurus huoyanshanensis" Dong 1977 into "Aublysodon huoyanshanensis". It probably represents a "Tarbosaurus" specimen.
History of paleontology
In 1808, Cuvier identified a fossil found in Maastricht as a giant marine reptile that would later be named "Mosasaurus". He also identified, from a drawing, another fossil found in Bavaria as a flying reptile and named it "Pterodactylus". He speculated, based on the strata in which these fossils were found, that large reptiles had lived prior to what he was calling "the age of mammals". Cuvier's speculation would be supported by a series of finds that would be made in Great Britain over the course of the next two decades. Mary Anning, a professional fossil collector since age eleven, collected the fossils of a number of marine reptiles from the Jurassic marine strata at Lyme Regis. These included the first ichthyosaur skeleton to be recognized as such, which was collected in 1811, and the first two plesiosaur skeletons ever found in 1821 and 1823. Many of her discoveries would be described scientifically by the geologists William Conybeare, Henry De la Beche, and William Buckland. It was Anning who observed that stony objects known as "bezoar stones" were often found in the abdominal region of ichthyosaur skeletons, and she noted that if such stones were broken open they often contained fossilized fish bones and scales as well as sometimes bones from small ichthyosaurs. This led her to suggest to Buckland that they were fossilized feces, which he named coprolites, and which he used to better understand ancient food chains.
Paranthodon
Paranthodon ( ) is a genus of stegosaurian dinosaur that lived in South Africa during the Early Cretaceous, between 139 and 131 million years ago. Discovered in 1845, it was one of the first stegosaurians found. Its only remains, a partial skull and isolated teeth, were found in the Kirkwood Formation. British paleontologist Richard Owen initially identified the fragments as those of the pareiasaur "Anthodon". After remaining untouched for years in the British Museum of Natural History, the partial skull was identified by South African paleontologist Robert Broom as belonging to a different genus; he named the specimen "Palaeoscincus africanus". Several years later, Hungarian paleontologist Franz Nopcsa, unaware of Broom's new name, similarly concluded that it represented a new taxon, and named it "Paranthodon owenii". Since Nopcsa's species name was assigned after Broom's, and Broom did not assign a new genus, both names are now synonyms of the current binomial, "Paranthodon africanus". The genus name combines the Ancient Greek "para" (near) with the genus name "Anthodon", to represent the initial referral of the remains.
Paleontology in Massachusetts
Among the earliest major fossil discoveries in Massachusetts occurred during the spring of 1802. Pliny Moody had uncovered a piece of reddish sandstone with bird-like three toed footprints while ploughing on his father's farm in South Hadley. More fossil footprints were discovered a few decades later, in 1835. At the time, Greenfield Massachusetts was paving when residents noticed footprints on the sandstone slabs that resembled those of turkeys. The rocks used in the project had been excavated at Turners Falls. This location would turn out to be the most productive dinosaur tracksite in the Connecticut Valley. Later that year, the fossil footprints discovered while paving Greenfield were brought to the attention of local naturalist and physician James Deane, whose curiosity was piqued. Deane wrote a letter to Amherst College geology professor Edward B. Hitchcock. Hitchcock spent the rest of the summer traveling through the Connecticut Valley examining the fossil footprints. The next year Hitchcock wrote a scientific paper on the fossil footprints of the Connecticut Valley. He thought the tracks were made by giant birds. Later, in 1843, James Deane began publishing on the Connecticut Valley track fossils with a paper in the American Journal of Science. In 1847 the ichnospecies Otozoum moodi was named after Pliny Moody.
Sauropoda
The first sauropod fossil to be scientifically described was a single tooth known by the non-Linnaean descriptor "Rutellum implicatum". This fossil was described by Edward Lhuyd in 1699, but was not recognized as a giant prehistoric reptile at the time. Dinosaurs would not be recognized as a group until over a century later.
Ichthyosaur
In 1811, in Lyme Regis, along what is now called the Jurassic Coast of Dorset, the first complete ichthyosaur skull was found by Joseph Anning, the brother of Mary Anning, who in 1812 while still a young girl, secured the torso of the same specimen. Their mother, Molly Anning, sold the combined piece to squire Henry Henley for £23. Henley lent the fossil to the London Museum of Natural History of William Bullock. When this museum was closed, the British Museum bought the fossil for a price of £47.5s; it still belongs to the collection of the now independent Natural History Museum and has the inventory number BMNH R.1158. It has been identified as a specimen of "Temnodontosaurus platyodon".
Dimorphodon
In 1858, Richard Owen reported the find of two new specimens, NHMUK PV R 41212 and NHMUK PV R 1035, again partial skeletons but this time including the skulls. Having found the skull to be very different from that of "Pterodactylus", Owen assigned "Pterodactylus macronyx" its own genus, which he named "Dimorphodon". His first report contained no description and the name remained a "nomen nudum". In 1859, however, a subsequent publication by Owen provided a description. After several studies highlighting aspects of "Dimorphodon" 's anatomy, Owen in 1874 made NHMUK PV R 1034 the holotype.
Feathered dinosaur
One of the earliest discoveries of possible feather impressions by non-avian dinosaurs is an ichnofossil ("Fulicopus lyellii") of the 195-199 million year old Portland Formation in the northeastern United States. Gierlinski (1996, 1997, 1998) and Kondrat (2004) have interpreted traces between two footprints in this fossil as feather impressions from the belly of a squatting dilophosaurid. Although some reviewers have raised questions about the naming and interpretation of this fossil, if correct, this early Jurassic fossil is the oldest known evidence of feathers, almost 30 million years older than the next-oldest-known evidence.
After a century of hypotheses without conclusive evidence, well-preserved fossils of feathered dinosaurs were discovered during the 1990s, and more continue to be found. The fossils were preserved in a Lagerstätte—a sedimentary deposit exhibiting remarkable richness and completeness in its fossils—in Liaoning, China. The area had repeatedly been smothered in volcanic ash produced by eruptions in Inner Mongolia 124 million years ago, during the Early Cretaceous epoch. The fine-grained ash preserved the living organisms that it buried in fine detail. The area was teeming with life, with millions of leaves, angiosperms (the oldest known), insects, fish, frogs, salamanders, mammals, turtles, and lizards discovered to date.
Pteranodon
Because the key distinguishing characteristic Marsh noted for "Pteranodon" was its lack of teeth, any toothless pterosaur jaw fragment, wherever it was found in the world, tended to be attributed to "Pteranodon" during the late nineteenth and early twentieth centuries. This resulted in a plethora of species and a great deal of confusion. The name became a wastebasket taxon, rather like the dinosaur "Megalosaurus", to label any pterosaur remains that could not be distinguished other than by the absence of teeth. Species (often dubious ones now known to be based on sexual variation or juvenile characters) have been reclassified a number of times, and several subgenera have in the 1970s been erected by Halsey Wilkinson Miller to hold them in various combinations, further confusing the taxonomy (subgenera include "Longicepia", "Occidentalia", and "Geosternbergia"). Notable authors who have discussed the various aspects of "Pteranodon" include Bennett, Padian, Unwin, Kellner, and Wellnhofer. Two species, "P. orogensis" and "P. orientalis", are not pteranodontids and have been renamed "Bennettazhia oregonensis" and "Bogolubovia orientalis" respectively.
Xenoposeidon
Xenoposeidon (meaning "strange or alien Poseidon", in allusion to "Sauroposeidon") is a genus of rebbachisaurid sauropod dinosaur from the Early Cretaceous of England, living about 140 million years ago. It is known from a single partial vertebra with unusual features, unlike those of other sauropods. This bone was first discovered in the early 1890s but received little attention until it was found by University of Portsmouth student Mike Taylor, who formally described and named it in 2007 with Darren Naish.
Pteranodon
"Pteranodon sternbergi" is the only known species of "Pteranodon" with an upright crest. The lower jaw of "P. sternbergi" was long. It was collected by George F. Sternberg in 1952 and described by John Christian Harksen in 1966, from the lower portion of the Niobrara Formation. It was older than "P. longiceps" and is considered by Bennett to be the direct ancestor of the later species.