92-702: Pteranodon ( / t ə ˈ r æ n ə d ɒ n / ; from Ancient Greek : πτερόν , romanized : pteron ' wing ' and ἀνόδων , anodon ' toothless ' ) is a genus of pterosaur that included some of the largest known flying reptiles , with P. longiceps having a wingspan of over 6 m (20 ft). They lived during the late Cretaceous geological period of North America in present-day Kansas , Nebraska , Wyoming , South Dakota and Alabama . More fossil specimens of Pteranodon have been found than any other pterosaur, with about 1,200 specimens known to science, many of them well preserved with nearly complete skulls and articulated skeletons. It
184-543: 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
276-423: A detailed description of the animal. 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. In a review of pterosaur size estimates published in 2010, researchers Mark Witton and Mike Habib argued that the largest estimate of 93 kg is much too high and an upper limit of 20 to 35 kg
368-486: A different type of toothless pterosaur. Williston was also the first scientist to critically evaluate all of the Pteranodon species classified by Cope and Marsh. He agreed with most of Marsh's classification, with a few exceptions. First, he did not believe that P. ingens and P. umbrosus could be considered synonyms, which even Cope had come to believe. He considered both P. velox and P. longiceps to be dubious;
460-467: A few males competing for association with groups consisting of large numbers of females. Similar to modern pinnipeds, Pteranodon may have competed to establish territory on rocky, offshore rookeries, with the largest, and largest-crested, males gaining the most territory and having more success mating with females. The crests of male Pteranodon would not have been used in competition, but rather as "visual dominance-rank symbols", with display rituals taking
552-540: A fossil collector working for Marsh. A second, smaller skull soon was discovered as well. These skulls showed that the North American pterosaurs were different from any European species, in that they lacked teeth and had bony crests on their skulls. Marsh recognized this major difference, describing the specimens as "distinguished from all previously known genera of the order Pterosauria by the entire absence of teeth." Marsh recognized that this characteristic warranted
644-477: 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
736-419: 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
828-453: A modern-day albatross . This is based on the fact that Pteranodon had a high aspect ratio (wingspan to chord length) similar to that of the albatross — 9:1 for Pteranodon , compared to 8:1 for an albatross. Albatrosses spend long stretches of time at sea fishing, and use a flight pattern called " dynamic soaring " which exploits the vertical gradient of wind speed near the ocean surface to travel long distances without flapping, and without
920-413: A more recent study suggests that it relied on thermal soaring, unlike modern seabirds but much like modern continental flyers and the extinct Pelagornis . Like other pterosaurs, Pteranodon probably took off from a standing, quadrupedal position. Using their long forelimbs for leverage, they would have vaulted themselves into the air in a rapid leap. Almost all of the energy would have been generated by
1012-420: A muscle attachment point. He suggested that the crest might have anchored large, long jaw muscles, but admitted that this function alone could not explain the large size of some crests. Bennett (1992) agreed with Eaton's own assessment that the crest was too large and variable to have been a muscle attachment site. Eaton had suggested that a secondary function of the crest might have been as a counterbalance against
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#17327806272041104-421: A new genus, and he coined the name Pteranodon ("wing without tooth") in 1876. Marsh reclassified all the previously named North American species from Pterodactylus to Pteranodon . He considered the smaller skull to belong to Pteranodon occidentalis , based on its size. Marsh classified the larger skull, YPM 1117, in the new species Pteranodon longiceps , which he thought to be a medium-sized species in between
1196-550: 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
1288-608: 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
1380-418: A single evolutionary lineage lasting about 4 million years. In other words, only one species of Pteranodon would have been present at any one time, and P. sternbergi (or Geosternbergia ) in all likelihood was the direct ancestor species of P. longiceps . Many researchers consider there to be at least two species of Pteranodon . However, aside from the differences between males and females described above,
1472-630: 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
1564-510: 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
1656-570: 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
1748-418: 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
1840-666: 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
1932-448: 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
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#17327806272042024-719: 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. In the early 1990s, Bennett noted that the two major morphs of pteranodont present in the Niobrara Formation were precisely separated in time with little, if any, overlap. Due to this, and to their gross overall similarity, he suggested that they probably represent chronospecies within
2116-453: Is more likely that Pteranodon could take off from the water, and would have dipped for fish while swimming rather than while flying. Even a small, female Pteranodon could have reached a depth of at least 80 centimeters (31 in) with its long bill and neck while floating on the surface, and they may have reached even greater depths by plunge-diving into the water from the air like some modern long-winged seabirds. In 1994, Bennett noted that
2208-471: Is more realistic. 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. Other distinguishing characteristics that set Pteranodon apart from other pterosaurs include narrow neural spines on
2300-413: 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 maiseyi in 2010,
2392-759: 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
2484-771: 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
2576-501: 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)
2668-1031: 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"):
2760-495: 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
2852-687: 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
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2944-659: The Niobrara Chalk, suggesting that animals who died on shore must have been washed out to sea (one specimen of a hadrosaur appears to have been scavenged by a shark ). 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 Santonian stage of the Cretaceous period. The genus
3036-508: The aid of thermals (which do not occur over the open ocean the same way they do over land). While most of a Pteranodon flight would have depended on soaring, like long-winged seabirds, it probably required an occasional active, rapid burst of flapping, and studies of Pteranodon wing loading (the strength of the wings vs. the weight of the body) indicate that they were capable of substantial flapping flight, contrary to some earlier suggestions that they were so big they could only glide. However,
3128-411: The angle of certain skull bones. Because well-preserved Pteranodon skull fossils are extremely rare, researchers use stratigraphy (i.e. which rock layer of the geologic formation a fossil is found in) to determine species identity in most cases. Pteranodon sternbergi is the only known species of Pteranodon with an upright crest. The lower jaw of P. sternbergi was 1.25 meters (4.1 ft) long. It
3220-550: 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
3312-419: 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
3404-468: The base of the jaws. The beaks were long, slender, and ended in thin, sharp points. The upper jaw, which was longer than the lower jaw, was curved upward; while this normally has been attributed only to the upward-curving beak, one specimen (UALVP 24238) has a curvature corresponding with the beak widening towards the tip. While the tip of the beak is not known in this specimen, the level of curvature suggests it would have been extremely long. The unique form of
3496-412: The beak in this specimen led Alexander Kellner to assign it to a distinct genus, Dawndraco , in 2010. The most distinctive characteristic of Pteranodon is its cranial crest. These crests consisted of skull bones (frontals) projecting upward and backward from the skull. The size and shape of these crests varied due to a number of factors, including age, sex, and species. Male Pteranodon sternbergi ,
3588-438: 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
3680-555: The concept of Pteranodon to include Nyctosaurus as a fourth subgenus. Miller considered these to be an evolutionary progression, with the primitive Nyctosaurus , at the time thought to be crestless, giving rise to Occidentalia (with a small crest), which in turn gave rise to Pteranodon with its long backwards crest, finally leading to Geosternbergia with its large, upright crest. However, Miller made several mistakes in his study concerning which specimens Marsh had assigned to which species, and most scientists disregarded his work on
3772-622: The correlation between crest size and body size. There is no evidence of extra blood vessels in the crest for this purpose, however, and the large, membranous wings filled with blood vessels would have served that purpose much more effectively. With these hypotheses ruled out, the best-supported hypothesis for crest function seems to be as a sexual display. This is consistent with the size variation seen in fossil specimens, where females and juveniles have small crests and males large, elaborate, variable crests. Adult Pteranodon specimens may be divided into two distinct size classes, small and large, with
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3864-454: The crest may have supported a membrane of skin connecting the backward-pointing crest to the neck and back, increasing its surface area and effectiveness as a rudder. The rudder hypothesis, again, does not take into account females nor P. sternbergi , which had an upward-pointing, not backward-pointing crest. Bennett also found that, even in its capacity as a rudder, the crest would not provide nearly so much directional force as simply maneuvering
3956-471: The crests of females had no counterbalancing effect, and that the crests of male P. sternbergi would, by themselves, have a negative effect on the balance of the head. In fact, side to side movement of the crests would have required more, not less, neck musculature to control balance. In 1943, Dominik von Kripp suggested that the crest may have served as a rudder , an idea embraced by several later researchers. One researcher, Ross S. Stein, even suggested that
4048-615: 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
4140-526: The discovery of pterosaur trackways . The possibility of aquatic locomotion via swimming has been discussed briefly in several papers (Bennett 2001, 1994, and Bramwell & Whitfield 1974). 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
4232-464: 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 renamed his giant North American pterosaur Pterodactylus occidentalis , meaning "Western wing finger," in his 1872 description of
4324-417: The first was based on non-diagnostic fragments, and the second, though known from a complete skull, probably belonged to one of the other, previously-named species. In 1903, Williston revisited the question of Pteranodon classification, and revised his earlier conclusion that there were seven species down to just three. He considered both P. comptus and P. nanus to be specimens of Nyctosaurus , and divided
4416-431: The forelimbs. The upstroke of the wings would have occurred when the animal cleared the ground followed by a rapid down-stroke to generate additional lift and complete the launch into the air. 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
4508-673: 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
4600-452: The fossil record, allowing for detailed descriptions of their anatomy and analysis of their life history. Over 1,000 specimens have been identified, though fewer than half of them are complete enough to give researchers good anatomical information. Still, this is more fossils material than is known for any other pterosaur, and it includes both male and female specimens of various age groups and possibly species. Adult Pteranodon specimens from
4692-433: The genus Pteranodon . However, both Williston and Pleininger were incorrect, because unnoticed by both of them was the fact that, in 1891, Seeley himself had finally described and properly named Ornithostoma , assigning it to the species O. sedgwicki . In the 2010s, more research on the identity of Ornithostoma showed that it was probably not Pteranodon or even a close relative, but may in fact have been an azhdarchoid ,
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#17327806272044784-403: The head, neck, and shoulders of Pteranodon were as heavily built as diving birds, and suggested that they could dive by folding back their wings like the modern gannet . Pteranodon was notable for its skull crest, though the function of this crest has been a subject of debate. Most explanations have focused on the blade-like, backward pointed crest of male P. longiceps , however, and ignored
4876-561: 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
4968-476: 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
5060-435: The large European genus Ornithocheirus , though he misspelled the name (forgetting the 'e'). Cope's paper naming his species was published in 1872, just five days after Marsh's paper. This resulted in a dispute, fought in the published literature, over whose names had priority in what obviously were the same species. Cope conceded in 1875 that Marsh's names did have priority over his, but maintained that Pterodactylus umbrosus
5152-399: The large crests only developed in males when they reached their large, adult size, making the sex of immature specimens difficult to establish from partial remains. The fact that females appear to have outnumbered males two to one suggests that, as with modern animals with size-related sexual dimorphism, such as sea lions and other pinnipeds , Pteranodon might have been polygynous , with
5244-477: The large size class being about one and a half times larger than the small class, and the small class being twice as common as the large class. Both size classes lived alongside each other, and while researchers had previously suggested that they represent different species, Christopher Bennett showed that the differences between them are consistent with the concept that they represent females and males, and that Pteranodon species were sexually dimorphic . Skulls from
5336-436: The larger size class preserve large, upward and backward pointing crests, while the crests of the smaller size class are small and triangular. Some larger skulls also show evidence of a second crest that extended long and low, toward the tip of the beak, which is not seen in smaller specimens. The gender of the different size classes was determined, not from the skulls, but from the pelvic bones. Contrary to what may be expected,
5428-674: 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 5.6 m (18 ft). Adult females were much smaller, averaging 3.8 m (12 ft) in wingspan. A large specimen of Pteranodon longiceps , USNM 50130, is estimated to have a wingspan of 6.25–6.5 m (20.5–21.3 ft), body length of 2.6 m (8.5 ft) and body mass of 50 kg (110 lb). Even larger specimens had wingspans of 7.25–7.6 m (23.8–24.9 ft). While most specimens are found crushed, enough fossils exist to put together
5520-411: The long beak, reducing the need for heavy neck muscles to control the orientation of the head. Wind tunnel tests showed that the crest did function as an effective counterbalance to a degree, but Bennett noted that, again, the hypothesis focuses only on the long crests of male P. longiceps , not on the larger crests of P. sternbergi and very small crests that existed among the females. Bennett found that
5612-457: The new species Pteranodon sternbergi , complicated the situation even further. prompting another revision of the genus by Miller in 1972. Because it was impossible to determine crest shape for all of the species based on headless skeletons, Miller concluded that all Pteranodon species except the two based on skulls ( P. longiceps and P. sternbergi ) must be considered nomena dubia and abandoned. The skull Eaton thought belonged to P. ingens
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#17327806272045704-441: The new specimen. He named two additional species, based on size differences: Pterodactylus ingens (the largest specimen so far), and Pterodactylus velox (the smallest). Meanwhile, Marsh's rival Edward Drinker Cope had unearthed several specimens of the large North American pterosaur. Based on these specimens, Cope named two new species, Ornithochirus umbrosus and Ornithochirus harpyia , in an attempt to assign them to
5796-508: 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
5888-457: The older species of the two described to date, had a more vertical crest with a broad forward projection, while their descendants, Pteranodon longiceps , evolved a narrower, more backward-projecting crest. Females of both species were smaller and bore small, rounded crests. The crests were probably mainly display structures, though they may have had other functions as well. The wing shape of Pteranodon suggests that it would have flown rather like
5980-418: The others into small ( P. velox ), medium ( P. occidentalis ), and large species ( P. ingens ), based primarily on the shape of their upper arm bones. He thought P. longiceps , the only one known from a skull, could be a synonym of either P. velox or P. occidentalis , based on its size. In 1910, Eaton became the first scientist to publish a more detailed description of the entire Pteranodon skeleton, as it
6072-487: 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
6164-479: The place of physical competition with other males. If this hypothesis is correct, it also is likely that male Pteranodon played little to no part in rearing the young; such a behavior is not found in the males of modern polygynous animals who father many offspring at the same time. Specimens assigned to Pteranodon have been found in both the Smoky Hill Chalk deposits of the Niobrara Formation , and
6256-415: The post-cranial skeletons of Pteranodon show little to no variation between species or specimens, and the bodies and wings of all pteranodonts were essentially identical. Two species of Pteranodon are traditionally recognized as valid: Pteranodon longiceps , the type species , and Pteranodon sternbergi . The species differ only in the shape of the crest in adult males (described above), and possibly in
6348-501: The pressures of fossilization, and concluded that no Pteranodon skeletons had any significant differences from each other besides their size. Therefore, Eaton was left to decide his classification scheme based on differences in the skulls alone, which he assigned to species just as Marsh did, by their size. In the end, Eaton recognized only three valid species: P. occidentalis , P. ingens , and P. longiceps . The discovery of specimens with upright crests, classified by Harksen in 1966 as
6440-570: 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 . It is likely that, as in other polygynous animals (in which males compete for association with harems of females), Pteranodon lived primarily on offshore rookeries, where they could nest away from land-based predators and feed far from shore; most Pteranodon fossils are found in locations which at
6532-470: 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
6624-427: The small P. occidentalis and the large P. ingens . Marsh also named several additional species: Pteranodon comptus and Pteranodon nanus were named for fragmentary skeletons of small individuals, while Pteranodon gracilis was based on a wing bone that he mistook for a pelvic bone. He soon realized his mistake, and re-classified that specimen again into a separate genus, which he named Nyctosaurus . P. nanus
6716-543: The smaller size class had disproportionately large and wide-set pelvic bones. Bennett interpreted this as indicating a more spacious birth canal, through which eggs would pass. He concluded that the small size class with small, triangular crests represent females, and the larger, large-crested specimens represent males. Note that the overall size and crest size also corresponds to age. Immature specimens are known from both females and males, and immature males often have small crests similar to adult females. Therefore, it seems that
6808-613: 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. Some very fragmentary fossils belonging to pteranodontian pterosaurs, and possibly Pteranodon itself, have also been found on
6900-460: The subgenus Longicepia , though this was later changed to simply Pteranodon due to the rules of priority. P. sternbergi and P. walkeri , the upright-crested species, were given the subgenus Sternbergia , which was later changed to Geosternbergia because Sternbergia was already in use ("preoccupied"). Finally, Miller named the subgenus Occidentalia for P. eatoni , the skull formerly associated with P. occidentalis . Miller further expanded
6992-641: The subject in their later research, though Wellnhofer (1978) followed Miller's species list. and Schoch (1984) somewhat oddly published another revision that essentially returned to Marsh's original classification scheme, most notably sinking P. longiceps as a synonym of P. ingens . During the early 1990s, S. Christopher Bennett also published several major papers reviewing the anatomy, taxonomy and life history of Pteranodon . Fragmentary fossils assigned to Pteranodon have also been discovered in Skåne , Sweden . Pteranodon species are extremely well represented in
7084-517: 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
7176-744: The time, were hundreds of kilometres from the coastline. Below the surface, the sea was populated primarily by invertebrates such as ammonites and squid . Vertebrate life, apart from basal fish, included sea turtles , such as Toxochelys , the plesiosaurs Elasmosaurus and Styxosaurus , and the flightless diving bird Parahesperornis . Mosasaurs were the most common marine reptiles, with genera including Clidastes , Mosasaurus and Tylosaurus . At least some of these marine reptiles are known to have fed on Pteranodon . Barnum Brown , in 1904, reported plesiosaur stomach contents containing "pterodactyl" bones, most likely from Pteranodon . Fossils from terrestrial dinosaurs also have been found in
7268-400: The torso, indicating that fish made up a majority of the diet of Pteranodon (though they may also have taken invertebrates). Traditionally, most researchers have suggested that Pteranodon would have taken fish by dipping their beaks into the water while in low, soaring flight. However, this was probably based on the assumption that the animals could not take off from the water surface. It
7360-493: The two major species can be divided into two distinct size classes. The smaller class of specimens have small, rounded head crests and very wide pelvic canals, even wider than those of the much larger size class. The size of the pelvic canal probably allowed the laying of eggs, indicating that these smaller adults are females. The larger size class, representing male individuals, have narrow hips and very large crests, which were probably for display. Adult male Pteranodon were among
7452-630: The upper layers, a fossil lacking the skull can be identified based on its position in the geologic column (though for many early fossil finds, precise data about its location was not recorded, rendering many fossils unidentifiable). Below is a cladogram showing the phylogenetic placement of this genus within Pteranodontia from Andres and Myers (2013). Muzquizopteryx coahuilensis "Nyctosaurus" lamegoi Nyctosaurus gracilis Ancient Greek language Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
7544-517: 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 25 centimeters (9.8 in) in the largest males. Unlike earlier pterosaurs, such as Rhamphorhynchus and Pterodactylus , Pteranodon had toothless beaks , similar to those of birds . Pteranodon beaks were made of solid, bony margins that projected from
7636-419: The wide range of variation across age and sex. The fact that the crests vary so much rules out most practical functions other than for use in mating displays. Therefore, display was probably the main function of the crest, and any other functions were secondary. Scientific interpretations of the crest's function began in 1910, when George Francis Eaton proposed two possibilities: an aerodynamic counterbalance and
7728-552: The wings. The suggestion that the crest was an air brake, and that the animals would turn their heads to the side in order to slow down, suffers from a similar problem. Additionally, the rudder and air brake hypotheses do not explain why such large variation exists in crest size even among adults. Alexander Kellner suggested that the large crests of the pterosaur Tapejara , as well as other species, might be used for heat exchange, allowing these pterosaurs to absorb or shed heat and regulate body temperature, which also would account for
7820-480: 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 ,
7912-619: Was a distinct species (but not genus) from any that Marsh had named previously. Re-evaluation by later scientists has supported Marsh's case, refuting Cope's assertion that P. umbrosus represented a larger, distinct species. While the first Pteranodon wing bones were collected by Marsh and Cope in the early 1870s, the first Pteranodon skull was found on May 2, 1876, along the Smoky Hill River in Wallace County (now Logan County), Kansas, USA, by Samuel Wendell Williston ,
8004-487: Was also later recognized as a Nyctosaurus specimen. In 1892, Samuel Williston examined the question of Pteranodon classification. He noticed that, in 1871, Seeley had mentioned the existence of a partial set of toothless pterosaur jaws from the Cambridge Greensand of England , which he named Ornithostoma . Because the primary characteristic Marsh had used to separate Pteranodon from other pterosaurs
8096-509: Was an important part of the animal community in the Western Interior Seaway . Pteranodon is the most famous pterosaur, frequently featured in dinosaur media and strongly associated with dinosaurs by the general public. While not dinosaurs, pterosaurs such as Pteranodon form a clade closely related to dinosaurs as both fall within the clade Avemetatarsalia . Pteranodon was the first pterosaur found outside of Europe . Its fossils first were found by Othniel Charles Marsh in 1871, in
8188-416: 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. Because fossils identifiable as P. sternbergi are found exclusively in the lower layers of the Niobrara Formation, and P. longiceps fossils exclusively in
8280-432: Was its lack of teeth, Williston concluded that "Ornithostoma" must be considered the senior synonym of Pteranodon . However, in 1901, Pleininger pointed out that "Ornithostoma" had never been scientifically described or even assigned a species name until Williston's work, and therefore had been a nomen nudum and could not beat out Pteranodon for naming priority. Williston accepted this conclusion and went back to calling
8372-445: Was known at the time. He used his findings to revise the classification of the genus once again based on a better understanding of the differences in pteranodont anatomy. Eaton conducted experiments using clay models of bones to help determine the effects of crushing and flattening on the shapes of the arm bones Williston had used in his own classification. Eaton found that most of the differences in bone shapes could be easily explained by
8464-433: Was placed in the new species Pteranodon marshi , and the skull Eaton assigned to P. occidentalis was re-named Pteranodon eatoni . Miller also recognized another species based on a skull with a crest similar to that of P. sternbergi ; Miller named this Pteranodon walkeri . To help bring order to this tangle of names, Miller created three categories or "subgenera" for them. P. marshi and P. longiceps were placed in
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