Instruct: Given a question, retrieve passages that answer the question.
Query: When were fossils of the Pararhabdodon first found?
Pararhabdodon has been classified in a number of different positions within Iguanodontia since its remains were first discovered in the mid-1980s.[1][11] When the first speciemens were originally discovered and described by Casanovas-Cladellas and colleagues, they were not thought to belong a new taxon at all, but merely to belong to the genus Rhabdodon, and of an uncertain species.[3] Not long after, they would be recognized of those of a unique species, and in 1993 they were given their modern name. At this point it was still thought to be a rhabdodontid or some other type of primitive iguanodont closely related to them. This was based on the characteristics of its vertebrae, as the cranial remains had not yet been found.[4] In 1994 these cranial remains would be found - two maxilla - and these would be used by the original authors to establish it as a species of hadrosaur.[6]
The earth formed around 4.6 billion years ago, with unicellular life emerging somewhat later after the cessation of the Late Heavy Bombardment, a period of intense asteroid impacts possibly caused by migration of the gas giant planets to their current orbits, however multicellularity and bodies are a relatively recent event in Earth’s history. Bodies first started appearing towards the end of the Precambrian Era, during the Ediacaran period. The fossils of the Ediacaran period were first found in Southern Australia in the Ediacara Hills, hence the name. However these fossils were initially thought to be part of the Cambrian and it wasn’t until the late 1950s when Martin Glaessner identified the fossils as actually being from the Precambrian era. The fossils that were found date to about 600 million years ago and are found in a variety of morphologies.
Pararhabdodon most likely was a bipedal-quadrupedal herbivore, in the neighborhood of 6m (19.7ft) long, fully grown. The dorsal and sacral neural spines were elongate, so the animal would have had a tall back, like other hadrosaurids. Because the material is sparse, more specific conclusions cannot be reached.[1]
Tsintaosaurini is a tribe of basal lambeosaurine hadrosaurs native to Eurasia. It currently contains only "Tsintaosaurus" (from Shandong, east China) and "Pararhabdodon" (from Spain and France).
"Koutalisaurus", also known from late Cretaceous Spain and formerly referred to "Pararhabdodon" 
, may also be a tsintaosaurin because of its association with the latter genus; some recent work also suggests it may indeed be referrable to "Pararhabdodon".
By 1949, when administration of the area passed to the Chinese Communist Party and its leader Mao Zedong, the fossils of Yixian were studied only by Chinese scientists. It was not until the 1990s when remarkable fossils of birds and dinosaurs were excavated. Since 1996, a number of dinosaur fossils that have revolutionized knowledge of these animals have been found at Yixian; among them are the first known non-avian theropods with feathers. See Jehol Biota.
Due to a well global distribution the "Ptychodus" is well represented in the fossil history; many fossils have been uncovered such as isolated teeth, fragments of dentition, calcified vertebral centra, denticles, and associated fragments of calcified cartilage. The very first remains of "Ptychodus" were found in England in the early 19th century and was characterized as "palates of fish".
The first discovery of Ptychodus teeth in Kansas came in 1868 when Leidy reported and described a damaged tooth near Fort Hays, Kansas. After, many more teeth were uncovered in almost perfect conditions and other species within the genus were identified. Fossils of species within this genus have been found in the marine strata of United States, Brazil, Canada, the Czech Republic, France, Germany, India, Japan, Jordan, Mexico, Sweden and the United Kingdom. The fact that so many fossils of "Ptychodus" have been found in different regions of the world provides evidence of a distribution of species during the Albian-Turonian time.
Anning searched for fossils in the area's Blue Lias cliffs, particularly during the winter months when landslides exposed new fossils that had to be collected quickly before they were lost to the sea. She nearly died in 1833 during a landslide that killed her dog, Tray. Her discoveries included the first ichthyosaur skeleton correctly identified; the first two more complete plesiosaur skeletons found; the first pterosaur skeleton located outside Germany; and important fish fossils. Her observations played a key role in the discovery that coprolites, known as bezoar stones at the time, were fossilised faeces. She also discovered that belemnite fossils contained fossilised ink sacs like those of modern cephalopods. When geologist Henry De la Beche painted "Duria Antiquior", the first widely circulated pictorial representation of a scene from prehistoric life derived from fossil reconstructions, he based it largely on fossils Anning had found, and sold prints of it for her benefit.
Evidence for this synonymy would later come in a 2009 study, from Prieto-Márquez alongside Jonathan R. Wagner. Material from Pararhabdodon, the holotype of Koutalisaurus, and material of the Chinese species Tsintaosaurus spinorhinus were examined and compared, and the edentulous slope previously thought unique to Koutalisaurus was found to be nearly identical in T. spinorhinus. It was noted various approaches could be taken to this matter. As the species K. koelerororum shared a unique trait with Tsintaosaurus and could not be distinguished from it, the former could be considered a junior synonym of the latter. However, it was noted that the likelihood of a large archosaur species with a large enough range to include both specimens is completely unprecedented, so they refrained from this referral. K. kohlerorum could instead be considered a nomen dubium, a taxon of uncertain distinctiveness due to lacking any autapomorphies. But they disliked this course of action too, as unless a second taxon with the same morphology was found in the area, they posited the specimen likely did represent a unique component of the local fauna. Thus, they decided to keep it distinct, relative to Tsintaosaurus. No material had, as yet, been discovered from the SRA locality permitting comparison with Pararhabdodon. However, traits uniting P. isonensis with T. spinorhinus were found. As both taxa from the Tremp Formation uniquely shared traits with the Asian genus, the authors decided to treat the two as one species, as maintaining them as provisionally separate would in their eyes be misleading to non-specialists, who would likely not distinguish the two taxa were kept separate to be conservative and not due to strong evidence for two hadrosaurs in the area.[11]
First of its fossils were cranial remains found in Portugal, and later more fossils were found in France and Spain. They are only known from very fragmentary fossils, elements of the skull, dentary, teeth and osteoderm.
Ancient people living near Fossil, Oregon collected fossils as far back as 11,000 years ago and kept them at a dwelling that has since been uncovered by archeologists. Some of the fossils kept there were pierced to be made into jewelry. Five slabs of rock bearing leaf impressions were found neatly stacked in the corner of the site. These and other fossils discovered by Native Americans may have contributed to the development of local myths and lore. Historian Adrienne Mayor cites as an example the Klamath Tribes of the Modoc, who attributed local fossils to water monsters killed by the mythological figure Coyote.
Oregon's first paleontologist was Thomas Condon. Condon began collecting in 1861, when a company of soldiers that arrived in Fort Dalles, where Condon served as the Congregational church pastor. The soldiers brought fossil bones and teeth, including a well-preserved rhinoceros jaw, from the John Day fossil beds. When soldiers were dispatched the following year to Harney Valley, near the fossil beds, Condon went along with them and prospected for fossils. He went fossil collecting again in 1863 and discovered rich fossil deposits north of Picture Gorge in the John Day River valley.
The Fossil Grove is on a by floor of an old quarry, and belongs to the same geological time period as several other groups of "Lepidodendron" fossils found northwest of Glasgow. The shales and sandstones exposed around the fossils belong to the Limestone Coal Formation of the Clackmannan Group, with the shale containing imprints of various plant remains. The strata were deposited approximately 325 million years ago when the trees were still alive, and the strata are now folded and cut by many faults and slant downwards to the northeast. Beds of coal and some ironstones can be found in the northern and eastern parts of the Grove. The quarry in which the fossils were found is an igneous dolerite sill that was intruded into the sediments and two of the trunks during the Early Permian period approximately 290 million years ago. The quarry was already mostly excavated when the fossils were found, with the dolerite being used to macadamize nearby roads.
The same study also evaluated Blasisaurus canudoi, from the Blasi 1 locality of the underlying Arén Formation, and Arenysaurus ardevoli, from the Blasi 3 locality of the earlier "gray unit" of the Tremp Formation. Previously claimed diagnostic traits of the two taxa were shown to be found in other hadrosaurs. This left the former with two unique characteristics - both in the jugal - and a unique combination of other characteristics, and the latter with only one unique characteristic - in the frontal. As overlapping fossils were not known for the relevant areas, it couldn't be ruled out that they were indeed synonymous, representatives of a single species. The same re-evaluation could not rule out that this single species or either of the separated species were synonyms of Pararhabdodon isonensis, as no taxonomically informative areas of the skeleton are known from both Pararhabdodon and the other two taxa. The authors refrained from considering any of them representatives of one species pending data from more material.[8]
Pararhabdodon (meaning "near fluted tooth" in reference to Rhabdodon) is a genus of tsintaosaurin hadrosaurid dinosaur, from the Maastrichtian-age Upper Cretaceous Tremp Formation of Spain. The first remains were discovered from the Sant Romà d’Abella fossil locality and assigned to the genus Rhabdodon, and later named as the distinct species Pararhabdodon isonensis in 1993. Known material includes assorted postcranial remains, mostly vertebrae, as well as maxillae from the skull. Specimens from other sites, including remains from France, a maxilla previously considered the distinct taxon Koutalisaurus kohlerorum, an additional maxilla from another locality, the material assigned to the genera Blasisaurus and Arenysaurus, and the extensive Basturs Poble bonebed have been considered at different times to belong to the species, but all of these assignments have more recently been questioned.
When the first fossils were discovered, they were mistakenly thought to be sponges, hence the generic name. Later, similarly shaped fossils were found, and (correctly) identified as those of a crinoid, and named ""Dinocrinus"." ("Terrible Lily") It was soon realized that "Dinocrinus" and "Calceolispongia" were the same, and "Dinocrinus" is now regarded as a junior synonym.
Near the village Abella de la Conca, in the 1990s, palaeontologist Marc Boada discovered a new site in the Tremp Formation, bearing dinosaur fossils. From this site, later named Les Llaus (LL), a right dentary, specimen designation IPS SRA 27, was excavated.[1] In 1997, Casanovas-Cladellas and colleagues stated this dentary was discovered from SRA, the site where the original Pararhabdodon remains were found.[1][6] The following year, they described the specimen and referred it to P. isonensis, and again stated it was from the same stratigraphic level.[2] In 2006, the stratigraphy of the region was re-evaluated, and the dentary's location was corrected to the LL locality, 750 metres (2,460ft) away from and 9 metres (30ft) below the SRA locality.[1]
In 2013, a maxilla, specimen designation MCD 4919, was referred to P. isonensis. It possessed synapomorphies of the tsintaosaurin tribe to which Pararhabdodon was by that point assigned to, and was referable to Pararhabdodon in particular due to rostrocaudally broad rostrocaudal region of the maxilla, a trait unique to P. isonensis relative to Tsintaosaurus, the other named tsintaosaurin taxon. The specimen was discovered at the Serrat del Rostiar 1 locality, part of the "lower red unit" of the Tremp formation just like the type locality, SRA, but in a lower section of the strata, making it older. This extended the range of species further within the upper Maastrichtian. The Serrat del Rostiar 1 locality is near the village of Basturns.[8] In 2019, the new genus and species Adynomosaurus arcanus was described, and in the paper describing the referral of MCD 4919 to P. isonensis was questioned. The specimen was found to have a differing ectopterygoid shelf to the maxilla of the SRA Pararhabdodon, which would conflict with their identity as belonging to the same species. As the specimen was still recognized as having tsintaosaurin characteristics, the authors still considered it to likely belong to a close relative of P. isonensis.[10]
Snakes do not have an extensive fossil record; the oldest known fossil is from between early and late Cretaceous period. There were Tertiary fossil snakes that became extinct by the end of the Eocene period. The first colubrid also appeared in the Eocene period. Lizards first appeared in the middle Jurassic period, and this is when the scincomorph and the anguimorph lizards were first seen. The Gekkotans first appear in the late Jurassic period and the iguanians first appear in the late Cretaceous period. The lizards of the Cretaceous period represent extinct genera and species. The majority of amphisbaenians first appeared during the early Cenozoic period. Rhynchocephalian fossils first appear in the Middle Triassic period, between 238-240 million years ago, making them the earliest lepidosaurian fossils found to date. The tuatara can now only be found on small islands off the New Zealand coast. However, fossil records show that it once lived on mainland New Zealand and Rhynchocephalia as a whole was once distributed globally.
Prieto-Márquez would return to the issue in 2009 along with Jonathan R. Wagner. They once again turned to the articulation between the maxilla and the jugal, finding this to link Pararhabdodon to the Asian lambeosaurine, Tsintaosaurus spinorhinus. In the 2006 phylogenetic analysis, P. isonenis had been scored as having an ancestral hadrosauroid condition for this trait, which had a strong effect on its position. Upon instead coding it for a modified version of the more advanced hadrosaurid condition, the connection between the two species to the exclusion of other lambeosaurines was supported. Multiple synapomorphies were found uniting to the two taxa, as well as identifying Pararhabdodon as a member of the subfamily as opposed to outside of Hadrosauridae.[11] This relationship was again supported in Prieto-Márquez et al. (2013), where the group containing the two genera was coined as the taxonomic tribe Tsintaosaurini; a diagnosis for the tribe was provided. Their cladogram is reproduced above, on the right.[8]
The proposal that P. isonensis was the first known European lambeosaurine was soon challenged, however. In 2001, Jason Head, in a study re-evaluating the status of another species, Eolambia caroljonesa, as a primitive lambeosaur. Both were determined to instead be more primitive hadrosauroids more basal than the split between Lambeosaurinae and "Hadrosaurinae" (later renamed to Saurolophinae[18]). In regards to Pararhabdodon, this was based on rebuttal of arguments put forward in the 1999 study; the present anatomy maxilla-jugal articulation was not considered a synapomorphy in Lambeosaurinae, merely the presence of this in the jugal itself, unknown in the species, and the angular deltopectoral crest is a trait present even in primitive iguanodontians. Additionally, its tooth count was lower than those of known lambeosaurs.[19] In the 2006 re-evaluation of the genus by Albert Prieto-Márquez and colleagues, it was included in a phylogenetic analysis for the first time. This found it to be a non-hadrosaurid hadrosauroid, a similar position as had been argued by Head. The cladogram of Prieto-Márquez et al. (2006) is seen below on the left:[1]
Elgin Reptiles is the name given to the Permian and Triassic fossils found in the sandstone deposits in and around the town of Elgin, in Moray, Scotland. They are of great historical and scientific importance, and many of the specimens are housed in the Elgin Museum, and some in the Hunterian in Glasgow, and the National Museum of Scotland in Edinburgh. The Elgin Reptiles include the dinosauriform "Saltopus elginensis", the dicynodont "Gordonia", and the pareiasaur "Elginia". There are also many footprints and tail-drags associated with the same Permian and Triassic sandstone deposits.
The sandstone in the Elgin area was originally quarried for building materials. The quarries were where the first reptile fossils were found, and they have continued to yield fossils to this day.
The first Elgin Reptile was discovered in 1844, but because it was only a few scales scientists of the time believed it was an Old Red Sandstone fish fossil, which were relatively well-known from sites across Northern Scotland. In 1851 a second indisputably reptilian fossil was found. Fossils of "Gordonia" were first found by Scottish naturalist Ramsay Heatley Traquair in 1885 and E.T. Newton described several key fossil specimens.
Prieto-Márquez returned again to the dentary in 2013, in a study alongside colleagues providing a review and investigation of hadrosaurs from all over Europe. Further preparation of the specimen in the time since his last study regarding it revealed the uniqueness of the dentary had been exaggerated significantly by reconstruction of the specimen when it was first prepared in the 1990s. The Tsintaosaurus specimens showing the similar condition were found to have been distorted from a similar process. Correcting for the innaccuracies, the Los Llaus dentary is indistinguishable from that of multiple lambeosaurines, and shares no particular connection to Tsintaosaurus. With this, their reasoning for assignment of the specimen to Pararhabdodon was voided, and the specimen is now considered a completely indeterminate lambeosaurine dentary.[8]
Fondevilla et al. (2018) compared to the material from Basturs Poble to recognized species of lambeosaurine from geographically and temporally similar locations in the Pyrenees. The jugals from the sample were found to be distinct from those known from Blasisaurus and Arenysaurus (possible synonyms of Pararhabdodon), and the frontals were also distinguishable from the latter. Identity as specimens as either of those taxa was therefore ruled out. Overlapping taxonomically informative material with Pararhabdodon was not found from the bonebed, excepting a maxilla too fragmentary for use in referral. However, both the type locality of SRA and the locality of the referred maxilla MCD 4919 are close to the area the bonebed was found. As such, it was considered most likely the Basturs Poble hadrosaurs are members of the species P. isonensis, but the poor state of preservation of the only comparable material in the bonebed prevented a secure referral.[12] Later, a 2019 study by Prieto-Márquez et al. described a new genus and species of Spanish lambeosaurine, Adynomosaurus arcanus. They compared the material of their species to that of the bonebed, and though similarities were found, their tooth crowns and ilia had different shapes, and the scapulae from the site lacked the distinct shallow nature that characterizes A. arcanus. Accordingly, they considered it unlikely the bonebed belonged to their species. Additionally, they commented on the lack of tsintaosaurin synapomorphies in the bonebed material and so kept a referral as indeterminate lambeosaurines rather than considering it to represent Pararhabdodon.[10]
Iguanodon bernissartensis Iguanodon atherfieldensis Probactrosaurus gobiensis Ouranosaurus nigeriensis Protohadros byrdi Bactrosaurus johnsoni Gilmoreosaurus mongoliensis Telmatosaurus transsylvanicus Tanius sinensis Pararhabdodon isonensisHadrosauridae Hadrosaurinae Lambeosaurinae Equijubus normani Probactrosaurus gobiensis Protohadros byrdi Eolambia caroljonesa Tanius sinensis Bactrosaurus johnsoni Telmatosaurus transsylvanicus Tethyshadros insularis Lophorhothon atopusHadrosauridae Hadrosaurus foulkii Edmontosaurus annectens Brachylophosaurus canadensisLambeosaurinae Aralosaurini Jaxartosaurus aralensis Tsintaosaurini Tsintaosaurus spinorhinus Pararhabdodon isonensis Parasaurolophini Lambeosaurini
Marc Boada discovered, in the late 1990s, a new fossil-bearing locality 300 metres (980ft) from the village of Basturs. As the village already lent its name to a locality where dinosaur eggs had been found, it was named the Basturs Poble (BP) locality, "poble" derived from the word for village. The locality belongs to the Conques Formation of the Tremp Group, dated to the late-Early Maastrictian.[12] Over ten years, from 2001 through 2011, field work was done at this site, and a massive bonebed - consisting of some one thousand skeletal elements - was discovered.[12][13] The discovery was first reported in the literature in an abstract at the "55th Symposium for Vertebrate Palaeontology and Comparative Anatomy and the 16th meeting of the Symposium for Palaeontological Preparation and Conservation" conference in 2007. The authors, then, approached the bonebed under the hypothesis of a single species being present at the site. It was reported that all ontogenetic stages were present.[14] Later, a poster presentation and abstract at another conference suggested the specimens may have belonged to Koutalisaurus kohlororum, but they cautioned an adult dentary from the bonebed would need to be discovered to test the hypothesis.[15] This was later rejected, as Koutalisaurus was found to be an invalid nomen dubium.[8] In 2018, the material was instead noted as possibly referrable to Pararhabdodon isonensis.[12]
"Tsintaosaurus" may form a clade in Lambeosaurinae with the European genera "Pararhabdodon" and "Koutalisaurus" (probable synonym of "Pararhabdodon").