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Ornithocheiromorpha

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74-560: Ornithocheiromorpha (from Ancient Greek , meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea . Fossil remains of this group date back from the Early to Late Cretaceous periods ( Valanginian to Turonian stages), around 140 to 92.5 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica , though most genera were recovered in Europe , Asia and South America . They were

148-543: A pitch accent . In Modern Greek, all vowels and consonants are short. Many vowels and diphthongs once pronounced distinctly are pronounced as /i/ ( iotacism ). Some of the stops and glides in diphthongs have become fricatives , and the pitch accent has changed to a stress accent . Many of the changes took place in the Koine Greek period. The writing system of Modern Greek, however, does not reflect all pronunciation changes. The examples below represent Attic Greek in

222-438: A wingspan ranging from 4.3 to 5 meters (14 to 16 ft), with the most complete known skull estimated to have been about 45 centimeters (1.48 ft) in length, based on a long-lost fragment of its jaw reported in 2012. Though its jaws measured only 28.5 centimeters (11.2 in), which was less than 80 percent of the skull's length. Anhanguerids and were typically larger than others of the group and were more successful within

296-490: A chalk pit near Burham in Kent , England . In 1846, British paleontologist James Scott Bowerbank named and described the remains found as Pterodactylus giganteus , as it was common at that time to assign any new described pterosaur species to Pterodactylus . In the same chalk pit as P. giganteus , two other pterosaur species were discovered. The first was named in 1851 by Bowerbank as Pterodactylus cuvieri , in honor of

370-493: A different clade than Pteranodon , though still within Pteranodontoidea . In 2003, David Unwin considered the family Istiodactylidae to group with the toothless Pteranodontidae, within the group Ornithocheiroidea , but Alexander Kellner however, grouped it with the toothed Anhangueridae instead, resulting in a more understandable change of evolution between the two toothed families. Brian Andres and colleagues found

444-419: A different genus called Nicorhynchus . Most anhanguerids bore distinctive convex "keeled" crests on their snout and underside of their mandible , this was well developed in several genera such as Tropeognathus . The similar Anhanguera possessed jaws that were tapered in width, but expanded into a broad, spoon-shaped rosette at the tip. The jaws are distinguished from its relatives by several differences in

518-471: A distinctive Brazilian school of the study of pterosaurs, with its own favoured phylogenetic model, clade terminology and nomenclature. Rival models and nomenclatural choices have been devised by the influential British pterosaur researcher David Unwin . Kellner has received numerous honours and prizes, including the TWAS Prize for Earth Sciences from The World Academy of Sciences . He was admitted in

592-477: A lack of contemporaneous evidence. Several theories exist about what Hellenic dialect groups may have existed between the divergence of early Greek-like speech from the common Proto-Indo-European language and the Classical period. They have the same general outline but differ in some of the detail. The only attested dialect from this period is Mycenaean Greek , but its relationship to the historical dialects and

666-419: A lesser degree. Pamphylian Greek , spoken in a small area on the southwestern coast of Anatolia and little preserved in inscriptions, may be either a fifth major dialect group, or it is Mycenaean Greek overlaid by Doric, with a non-Greek native influence. Regarding the speech of the ancient Macedonians diverse theories have been put forward, but the epigraphic activity and the archaeological discoveries in

740-550: A prefix /e-/, called the augment . This was probably originally a separate word, meaning something like "then", added because tenses in PIE had primarily aspectual meaning. The augment is added to the indicative of the aorist, imperfect, and pluperfect, but not to any of the other forms of the aorist (no other forms of the imperfect and pluperfect exist). The two kinds of augment in Greek are syllabic and quantitative. The syllabic augment

814-455: A rounded snout, very similar to that of Ornithocheirus and Anhanguera , and therefore it is placed within either Anhangueridae or Ornithocheiridae, depending on the author. Caulkicephalus was also a large pterosaur, with wingspan estimates of around 5 meters (16 ft). The vertebral column of ornithocheiromorphs was heavily pneumatized by an extensive system of air sacs, leaving prominent pneumatic foraminae. The neck of ornithocheiromorphs

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888-608: A separate historical stage, though its earliest form closely resembles Attic Greek , and its latest form approaches Medieval Greek . There were several regional dialects of Ancient Greek; Attic Greek developed into Koine. Ancient Greek was a pluricentric language , divided into many dialects. The main dialect groups are Attic and Ionic , Aeolic , Arcadocypriot , and Doric , many of them with several subdivisions. Some dialects are found in standardized literary forms in literature , while others are attested only in inscriptions. There are also several historical forms. Homeric Greek

962-630: A standard subject of study in educational institutions of the Western world since the Renaissance . This article primarily contains information about the Epic and Classical periods of the language, which are the best-attested periods and considered most typical of Ancient Greek. From the Hellenistic period ( c.  300 BC ), Ancient Greek was followed by Koine Greek , which is regarded as

1036-510: A vowel or /n s r/ ; final stops were lost, as in γάλα "milk", compared with γάλακτος "of milk" (genitive). Ancient Greek of the classical period also differed in both the inventory and distribution of original PIE phonemes due to numerous sound changes, notably the following: The pronunciation of Ancient Greek was very different from that of Modern Greek . Ancient Greek had long and short vowels ; many diphthongs ; double and single consonants; voiced, voiceless, and aspirated stops ; and

1110-434: A whole, similar to other ornithocheiroids . The three pelvic bones were often fused, as seen in many species such as Anhanguera santanae , the ilium was long and low, and its front and rear blades projected horizontally beyond the edges of the lower pelvic bones. Despite the structure length, the processes of these rod-like forms indicate that the hindlimb muscles attached to them were limited in strength. The pubic bone

1184-516: Is Guidraco . Its holotype skull has a length of 38 centimeters (15 in), which makes it smaller than other genera. The skull is very elongated however, and a hollow profile is seen, but not very pointed, as the upper edge and the line of the jaw run nearly parallel over most of their length. Even though most ornithocheiromorphs didn't have a cranial crest like the closely related pteranodontids, there were some exceptions, this included Caulkicephalus and Ludodactylus . Caulkicephalus had

1258-611: Is cladogram following a topology recovered by Brian Andres, using the most recent iteration of his data set (Andres, 2021). Pteranodontia [REDACTED] Serradraco Aussiedraco Hongshanopterus Targaryendraco Lonchodectes [REDACTED] Ikrandraco Lonchodraco Istiodactylidae Boreopteridae Hamipterus Brasileodactylus Barbosania Cearadactylus Ornithocheiridae Anhangueridae [REDACTED] Ludodactylus [REDACTED] Guidraco Cimoliopterus [REDACTED] Camposipterus Aetodactylus Below

1332-494: Is a cladogram showing the results of a phylogenetic analysis presented by Longrich et al. (2018). In the analysis, they placed the genus Hongshanopterus as a basal member. Hongshanopterus Lonchodraco Lonchodectes Boreopteridae Istiodactylidae Aetodactylus Cimoliopterus [REDACTED] Guidraco Ludodactylus [REDACTED] Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes

1406-480: Is a leading expert in the field of studying pterosaurs . His research has focused mainly on fossil reptiles from the Cretaceous Period , including extinct dinosaurs and crocodylomorphs . Kellner has over 500 publications to his name, has published more than 160 primary studies and two science books. He has participated in paleontological expeditions to many locations including Brazil, Chile, Iran,

1480-570: Is a literary form of Archaic Greek (derived primarily from Ionic and Aeolic) used in the epic poems , the Iliad and the Odyssey , and in later poems by other authors. Homeric Greek had significant differences in grammar and pronunciation from Classical Attic and other Classical-era dialects. The origins, early form and development of the Hellenic language family are not well understood because of

1554-418: Is added to stems beginning with consonants, and simply prefixes e (stems beginning with r , however, add er ). The quantitative augment is added to stems beginning with vowels, and involves lengthening the vowel: Some verbs augment irregularly; the most common variation is e → ei . The irregularity can be explained diachronically by the loss of s between vowels, or that of the letter w , which affected

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1628-666: Is called 'East Greek'. Arcadocypriot apparently descended more closely from the Mycenaean Greek of the Bronze Age. Boeotian Greek had come under a strong Northwest Greek influence, and can in some respects be considered a transitional dialect, as exemplified in the poems of the Boeotian poet Pindar who wrote in Doric with a small Aeolic admixture. Thessalian likewise had come under Northwest Greek influence, though to

1702-448: Is considered by some linguists to have been closely related to Greek . Among Indo-European branches with living descendants, Greek is often argued to have the closest genetic ties with Armenian (see also Graeco-Armenian ) and Indo-Iranian languages (see Graeco-Aryan ). Ancient Greek differs from Proto-Indo-European (PIE) and other Indo-European languages in certain ways. In phonotactics , ancient Greek words could end only in

1776-403: Is not well known in ornithocheiromorphs, however. Zhenyuanopterus , which is known for having 13 caudal vertebrae, formed one of the longest tails of any pterodactyloid. Anhanguera , another well-known genus, had a shorter tail, with broad caudal vertebrae that bore a "duplex" cross-section similar to Pteranodon . The pelvis of ornithocheiromorphs was of moderate size compared to the body as

1850-596: The Brazilian Congress . Since 2004 he has written a monthly column in Caçadores de Fósseis (Fossil Hunters), on the website Ciência Hoje On Line , a project of the Brazilian Society for Scientific Progress . Apart from studying their fossils, Kellner has performed important theoretical work on pterosaurs, including cladistic studies regarding their phylogeny . In this he is the founder of

1924-759: The Greek region of Macedonia during the last decades has brought to light documents, among which the first texts written in Macedonian , such as the Pella curse tablet , as Hatzopoulos and other scholars note. Based on the conclusions drawn by several studies and findings such as Pella curse tablet , Emilio Crespo and other scholars suggest that ancient Macedonian was a Northwest Doric dialect , which shares isoglosses with its neighboring Thessalian dialects spoken in northeastern Thessaly . Some have also suggested an Aeolic Greek classification. The Lesbian dialect

1998-671: The National Order of Scientific Merit (class Comendador), one the Brazilian Federal Government's most prestigious awards. Kellner's scientific achievements include the description of more than thirty species , of which Santanaraptor placidus (1996, 1999) is among the best examples of soft tissue preservation, including blood vessels and muscle fibers, reported in any dinosaur . The pterosaur Thalassodromeus sethi , which Kellner described in 2002 with his colleague Diogenes de Almeida Campos, allowed for

2072-642: The Paleontological Society of Chile . He is a research associate of the American Museum of Natural History and of the Chinese Institute of Vertebrate Paleontology and Paleoanthropology . Besides his teaching activity, having advised over fifteen master and Ph.D. students, Kellner has been active in the propagation of scientific knowledge to the general public. He organised the 1999 exposition No Tempo dos Dinossauros ("In

2146-507: The food chain rather than other ornithocheiromorphs, one reason is because of their large size, for example, Tropeognathus mesembrinus , had a normal wingspan of about 8.26 meters (27.1 ft), and 8.70 meters (28.5 ft) as the maximum estimate. Another species which was impressively large is Coloborhynchus capito , with a total skull length that could have been up to 75 centimeters (2.46 ft), leading to an estimated wingspan of 7 meters (23 ft). However, this species may belong to

2220-501: The present , future , and imperfect are imperfective in aspect; the aorist , present perfect , pluperfect and future perfect are perfective in aspect. Most tenses display all four moods and three voices, although there is no future subjunctive or imperative. Also, there is no imperfect subjunctive, optative or imperative. The infinitives and participles correspond to the finite combinations of tense, aspect, and voice. The indicative of past tenses adds (conceptually, at least)

2294-1031: The 5th century BC. Ancient pronunciation cannot be reconstructed with certainty, but Greek from the period is well documented, and there is little disagreement among linguists as to the general nature of the sounds that the letters represent. /oː/ raised to [uː] , probably by the 4th century BC. Greek, like all of the older Indo-European languages , is highly inflected. It is highly archaic in its preservation of Proto-Indo-European forms. In ancient Greek, nouns (including proper nouns) have five cases ( nominative , genitive , dative , accusative , and vocative ), three genders ( masculine , feminine , and neuter ), and three numbers (singular, dual , and plural ). Verbs have four moods ( indicative , imperative , subjunctive , and optative ) and three voices (active, middle, and passive ), as well as three persons (first, second, and third) and various other forms. Verbs are conjugated through seven combinations of tenses and aspect (generally simply called "tenses"):

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2368-500: The American Museum of Natural History in 1996. As well, he was involved in the organisation of several scientific meetings in Rio de Janeiro such as the 31st International Geological Congress in 2000 and the 2nd Latin-American Congress of Vertebrate Paleontology in 2002. Kellner has organized or taken part in several paleontological expeditions to many locations around the globe, including Brazil— Mato Grosso , Rio Grande do Sul and Ceará ;

2442-495: The Archaic period of ancient Greek (see Homeric Greek for more details): Μῆνιν ἄειδε, θεά, Πηληϊάδεω Ἀχιλῆος οὐλομένην, ἣ μυρί' Ἀχαιοῖς ἄλγε' ἔθηκε, πολλὰς δ' ἰφθίμους ψυχὰς Ἄϊδι προΐαψεν ἡρώων, αὐτοὺς δὲ ἑλώρια τεῦχε κύνεσσιν οἰωνοῖσί τε πᾶσι· Διὸς δ' ἐτελείετο βουλή· ἐξ οὗ δὴ τὰ πρῶτα διαστήτην ἐρίσαντε Ἀτρεΐδης τε ἄναξ ἀνδρῶν καὶ δῖος Ἀχιλλεύς. The beginning of Apology by Plato exemplifies Attic Greek from

2516-761: The Brazilian Sky") and the novel Na terra dos titãs ("In the Land of the Titans"). He has also taken part in documentaries about fossils (e.g., Antarctica - a Summer of 70 million Years and Dinosaur Hunters ). As a result of his scientific activity he has received several honours, being appointed a member of the Brazilian Academy of Sciences in 1997. He is also an honourable member of the New York Paleontologial Society and

2590-609: The Classical period of ancient Greek. (The second line is the IPA , the third is transliterated into the Latin alphabet using a modern version of the Erasmian scheme .) Ὅτι [hóti Hóti μὲν men mèn ὑμεῖς, hyːmêːs hūmeîs,   Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who

2664-618: The Time of the Dinosaurs") at the Museum of Earth Sciences , which has been regarded as a landmark for the establishment of paleontology in Brazil, attracting the attention of the people of Brazilian to the studies of fossils. In 2006 he organized the mounting of the first large-scale dinosaur skeleton in Brazil, that of the sauropod Maxakalisaurus topai , for which he received recognition from

2738-466: The United States, Argentina, China, and Antarctica. His scientific achievements include the description of more than thirty species . For his work he has received several honors and prizes, including the TWAS Prize for Earth Sciences from The World Academy of Sciences and admission to the National Order of Scientific Merit (class Comendador), one Brazil's most prestigious awards. Kellner

2812-574: The anhanguerids Tropeognathus and Anhanguera from the Romualdo Formation in Brazil . Tropeognathus was described with its type species, T. mesembrinus in 1987 by German paleontologist Peter Wellnhofer . The generic name is derived from Greek τρόπις, tropis , meaning "keel", and γνάθος, gnathos , meaning "jaw". The specific name is derived from Koine mesembrinos , "of the noontide", simplified as "southern", in reference to

2886-425: The anterior part of the head was upside down. Some of the teeth were extensively restored and enlarged until the wider front of the jaws showed very large and robust teeth projecting outward. With this arrangement, the maxilla was kinked, and its interlocking teeth suggested that Cearadactylus had a piscivourous diet, allowing the animal to keep hold of slippery fish. Another smaller genus similar to Cearadactylus

2960-550: The aorist. Following Homer 's practice, the augment is sometimes not made in poetry , especially epic poetry. The augment sometimes substitutes for reduplication; see below. Almost all forms of the perfect, pluperfect, and future perfect reduplicate the initial syllable of the verb stem. (A few irregular forms of perfect do not reduplicate, whereas a handful of irregular aorists reduplicate.) The three types of reduplication are: Irregular duplication can be understood diachronically. For example, lambanō (root lab ) has

3034-419: The augment when it was word-initial. In verbs with a preposition as a prefix, the augment is placed not at the start of the word, but between the preposition and the original verb. For example, προσ(-)βάλλω (I attack) goes to προσ έ βαλoν in the aorist. However compound verbs consisting of a prefix that is not a preposition retain the augment at the start of the word: αὐτο(-)μολῶ goes to ηὐ τομόλησα in

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3108-460: The belly ribs. The hip joint of ornithocheiromorphs was not perforated and allowed considerable mobility to the leg, and suggests that it was vertical, as therefore had a function in breathing, compensating the relative rigidity of the chest cavity. Several studies show that ornithocheiromorphs were less derived than the toothless pteranodontids such as Pteranodon , and based on the different evolutionary changes, they therefore need to be grouped in

3182-438: The center of Greek scholarship, this division of people and language is quite similar to the results of modern archaeological-linguistic investigation. One standard formulation for the dialects is: West vs. non-West Greek is the strongest-marked and earliest division, with non-West in subsets of Ionic-Attic (or Attic-Ionic) and Aeolic vs. Arcadocypriot, or Aeolic and Arcado-Cypriot vs. Ionic-Attic. Often non-West

3256-595: The chairman of these departments and has been head of the zoology graduate program since 2008. Kellner is also chief editor of the Anais da Academia Brasileira de Ciências , the official publication of the Brazilian Academy of Sciences . As part of his work on flying reptiles, Kellner organised the Pterosaur Workshop at Pittsburgh, USA, in 1995, and the first pterosaur symposium ever held at

3330-408: The crest and teeth: unlike its close relatives Coloborhynchus and Ornithocheirus , the crest on the upper jaw of Anhanguera didn't begin at the tip of the snout, therefore, it was set farther back on the skull. Other anhanguerids like Cearadactylus had its first preparations with many serious mistakes: the front of the snout and the lower jaws were confused leading to a reconstruction in which

3404-544: The deserts of Atacama , Chile and Kerman , Iran; Montana in the United States; Patagonia in Argentina; the famous deposits of Liaoning , China, and James Ross Island in Antarctica. Kellner has over 500 publications to his name (including abstracts and science articles). He has published more than 160 primary studies and two science books: Pterossauros - os senhores do céu do Brasil ("Pterosaurs — Lords of

3478-615: The dialect of Sparta ), and Northern Peloponnesus Doric (including Corinthian ). All the groups were represented by colonies beyond Greece proper as well, and these colonies generally developed local characteristics, often under the influence of settlers or neighbors speaking different Greek dialects. After the conquests of Alexander the Great in the late 4th century BC, a new international dialect known as Koine or Common Greek developed, largely based on Attic Greek , but with influence from other dialects. This dialect slowly replaced most of

3552-401: The families Istiodactylidae, Ornithocheiridae and Anhangueridae to form a group in 2014, which he called Lanceodontia, and consists of the more advanced ornithocheiromorphs. The clade however, excludes the more poorly known family Lonchodectidae, even though members of the family had previously been seen as some of the most derived forms of toothed pterosaurs. In their analysis, they also included

3626-516: The family Boreopteridae within the clade Anhangueria , though placed in a more basal position, while also containing the genus Guidraco . In 2018 however, Nicholas Longrich and colleagues found Boreopteridae outside Anhangueria as the sister taxon of the family Lonchodectidae, both groups placed as basal members of the Ornithocheiromorpha. There are competing theories of ornithocheiromorph phylogeny (evolutionary relationships). Below

3700-673: The forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek ( c.  1400–1200 BC ), Dark Ages ( c.  1200–800 BC ), the Archaic or Epic period ( c.  800–500 BC ), and the Classical period ( c.  500–300 BC ). Ancient Greek was the language of Homer and of fifth-century Athenian historians, playwrights, and philosophers . It has contributed many words to English vocabulary and has been

3774-502: The genus Ornithodesmus , as the vertebrae of the specimen of that genus was based on differed markedly from those of Hooley's specimen. In 1993, the British paleontologists Stafford C. Howse and Andrew C. Milner concluded that the holotype sacrum and only specimen of O. cluniculus didn't belong to a pterosaur, but instead to a maniraptoran theropod dinosaur. They also pointed out that no detailed attempts had been made to compare

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3848-778: The geologic Santana Group , about which he published many papers in the late 1980s. He earned a Master of Science degree in geology at the UFRJ in 1991, a MPhil degree in 1994, and a Ph.D. in 1996 from Columbia University in New York, in a joint program with the American Museum of Natural History . In 1997 he became a professor at the Federal University-owned National Museum of Rio de Janeiro and curator of its geological and paleontological departments. From 1998 to 2001, he served as

3922-561: The historical Dorians . The invasion is known to have displaced population to the later Attic-Ionic regions, who regarded themselves as descendants of the population displaced by or contending with the Dorians. The Greeks of this period believed there were three major divisions of all Greek people – Dorians, Aeolians, and Ionians (including Athenians), each with their own defining and distinctive dialects. Allowing for their oversight of Arcadian, an obscure mountain dialect, and Cypriot, far from

3996-476: The historical circumstances of the times imply that the overall groups already existed in some form. Scholars assume that major Ancient Greek period dialect groups developed not later than 1120 BC, at the time of the Dorian invasions —and that their first appearances as precise alphabetic writing began in the 8th century BC. The invasion would not be "Dorian" unless the invaders had some cultural relationship to

4070-458: The largest animals to have ever flown. Members of this group are also regarded to have some of the largest pterosaur wingspans , such as the one estimated for the huge Tropeognathus , though still not as large as those estimated for the azhdarchids , which may have reached up to 12 meters (39 ft). When ornithocheiromorphs first appeared, they were initially scavengers , consisting in a more terrestrial setting, but their success had made them

4144-533: The most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by pteranodontians and azhdarchoids . The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus , but not Pteranodon . Ornithocheiromorphs are considered to be some of

4218-508: The older dialects, although the Doric dialect has survived in the Tsakonian language , which is spoken in the region of modern Sparta. Doric has also passed down its aorist terminations into most verbs of Demotic Greek . By about the 6th century AD, the Koine had slowly metamorphosed into Medieval Greek . Phrygian is an extinct Indo-European language of West and Central Anatolia , which

4292-487: The perfect stem eilēpha (not * lelēpha ) because it was originally slambanō , with perfect seslēpha , becoming eilēpha through compensatory lengthening. Reduplication is also visible in the present tense stems of certain verbs. These stems add a syllable consisting of the root's initial consonant followed by i . A nasal stop appears after the reduplication in some verbs. The earliest extant examples of ancient Greek writing ( c.  1450 BC ) are in

4366-422: The prominent German naturalist and zoologist Georges Cuvier , while the second was described in the same year by British paleontologist Sir Richard Owen as Pterodactylus compressirostris . P. compressirostris later became the type species of a newly created genus called Lonchodectes (meaning " lance biter") in a review by English paleontologist Reginald Walter Hooley in 1914. Confusingly, this species

4440-750: The provenance from the Southern hemisphere . The description then led to an enormous taxonomic confusion. In 1989, Brazilian paleontologist Alexander Kellner considered it an Anhanguera mesembrinus , then a Coloborhynchus mesembrinus by Veldmeijer in 1998, and then a Criorhynchus mesembrinus in 2001 by German paleontologist Michael Fastnacht . T. mesembrinus was then considered a junior synonym of Ornithocheirus simus by British paleontologist David Unwin in 2001, but he then proposed an Ornithocheirus mesembrinus in 2003. In 2013 however, Taissa Rodrigues and Alexander Kellner concluded that Tropeognathus would be valid again, and containing only T. mesembrinus ,

4514-419: The sacrum of O. cluniculus with those of pterosaurs, and that O. latidens had in effect been treated as the type species of the genus Ornithodesmus . Howse, Milner, and David Martill in 2001, moved " O. " latidens to a new genus called Istiodactylus . They had also named a new family called Istiodactylidae , with Istiodactylus as the only member. Other important ornithocheiromorph discoveries include

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4588-436: The same site. He then considered it another species of Ornithodesmus . In 1901, Seeley named this new species as O. latidens , meaning "wide tooth". Later, Reginald Hooley discussed O. latidens in detail, based on specimens he had found, which led Ornithodesmus to be placed within a new family called Ornithodesmidae. Paleontologist Charles William Andrews however, had expressed doubts as to whether O. latidens belonged in

4662-552: The same study, Rodrigues & Kellner also reviewed the species Pterodactylus giganteus , and reassigned it to a newly created genus called Lonchodraco , this resulted in a new combination called Lonchodraco giganteus . In 1887, Seeley had described new fossil remains from the Isle of Wight , an island off the coast of southern England. He thought it belonged to some kind of bird-like creature, which he named it Ornithodesmus cluniculus . Seeley also reported another specimen found on

4736-517: The syllabic script Linear B . Beginning in the 8th century BC, however, the Greek alphabet became standard, albeit with some variation among dialects. Early texts are written in boustrophedon style, but left-to-right became standard during the classic period. Modern editions of ancient Greek texts are usually written with accents and breathing marks , interword spacing , modern punctuation , and sometimes mixed case , but these were all introduced later. The beginning of Homer 's Iliad exemplifies

4810-528: The time that pterosaurs were the ancestors of modern birds. In 1870, Seeley reassigned the species Pterodactylus cuvieri as Ornithocheirus cuvieri . In 1874, Richard Owen proposed two new genera, Coloborhynchus , meaning "maimed beak", and Criorhynchus , meaning "ram beak". While Coloborhynchus consisted in a totally new type species, C. clavirostris , as well as two other species reassigned from Ornithocheirus , Criorhynchus consisted entirely of former Ornithocheirus species, including O. simus , which

4884-570: The top predators of the skies, as well as the most common type of fish-eating pterosaur throughout the early Late Cretaceous. Some paleontologists also consider ornithocheiromorphs an earlier step of evolution to the pteranodontians , this is due to the similar flying techniques and flight locomotions, as well as their diet, which mainly consisted of fish, and therefore also hunted very similarly. Ornithocheiromorphs also flew like soaring birds, keeping their wings stretched and rarely flapping. The first specimens of ornithocheiromorphs were unearthed at

4958-500: The type species. A discovery in Asia, specifically northwestern China , was reported in 2006. The lake sediments allowed an exceptional preservation of fossils, and therefore paleontologists Qiu Zhanxiang and Wang Banyue started official excavations. Part of the findings consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. In 2014, a new species was named and described: Hamipterus tianshanensis . It

5032-480: Was Aeolic. For example, fragments of the works of the poet Sappho from the island of Lesbos are in Aeolian. Most of the dialect sub-groups listed above had further subdivisions, generally equivalent to a city-state and its surrounding territory, or to an island. Doric notably had several intermediate divisions as well, into Island Doric (including Cretan Doric ), Southern Peloponnesus Doric (including Laconian ,

5106-446: Was also long regarded, incorrectly, as the type species of Ornithocheirus . In 1861, further pterosaur specimens were found in the UK , and were given the new species Pterodactylus simus by Owen. British paleontologist Harry Govier Seeley then created the new genus Ornithocheirus for the new species in the same year, the generic name translating as "bird hand" is due to the notion of

5180-690: Was born in Vaduz , Liechtenstein , son of a German father and Austrian mother. In his early childhood he moved with his parents to Brazil, where he became naturalised Brazilian. In Rio de Janeiro he received a primary and secondary education at the bilingual Escola Alemã Corcovado . He began studying geology at the Federal University of Rio de Janeiro (UFRJ) in 1981. As a student, he soon began researching fossil vertebrates , particularly reptiles from Brazil's Cretaceous Period . His studies largely concerned pterosaur specimens from

5254-400: Was fused with the broad ischium into an ischiopubic blade, resulting in a narrow build. Sometimes, the blades of both sides were fused, closing the pelvis from below and forming the pelvic canal. The front of the pubic bones was also articulated with a unique structure, resulting in a pair of prepubic bones within. This formed a cusp covering the rear belly, and was located between the pelvis and

5328-443: Was later reassigned by Owen as Criorhynchus simus . In 2013, Brazilian paleontologists Taissa Rodrigues & Alexander Kellner made a deeper analysis on the species Pterodactylus cuvieri . In the analysis, they stated that it needed a separate genus, and assigning it to Ornithocheirus was inappropriate, therefore, they created the new genus called Cimoliopterus , with the new resulting combination Cimoliopterus cuvieri . In

5402-644: Was named by Wang Xiaolin, Alexander Kellner, Jiang Shunxing, Wang Qiang, Ma Yingxia, Yahefujiang Paidoula, Cheng Xin, Taissa Rodrigues, Meng Xi, Zhang Jialiang, Li Ning, and Zhou Zhonghe. The generic name Hamipterus combines that of the Hami region, with the word pteron , meaning "wing", and the specific name refers to the provenance from the Tian Shan , a mountain range. Ornithocheiromorphs were large pterosaurs, with wingspans normally ranging between 3 and 6 meters (9.8 and 19.7 ft). Istiodactylus for example, had

5476-410: Was typically relatively long and robust, being longer than the torso in some derived clades. The neural spines of ornithocheiromorph cervical vertebrae were generally tall and spikelike. In some genera such as Tropeognathus and Istiodactylus , up to six dorsal vertebrae are fused into a notarium . In some genera such as Anhanguera , four to seven sacral vertebrae are fused into a synsacrum . The tail

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