121-556: Thescelosaurus ( / ˌ θ ɛ s ɪ l ə ˈ s ɔː r ə s / THESS -il-ə- SOR -əs ; ancient Greek θέσκελος - ( theskelos - ) meaning "marvelous", and σαυρος ( sauros ) "lizard") is an extinct genus of neornithischian dinosaur that lived during the Late Cretaceous period in North America. It was among the last of the non-avian dinosaurs to appear before the entire group went extinct during
242-543: 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
363-580: A complete skull, most of a skeleton, and a mass in the chest cavity that was originally interpreted as a heart. Another, Museum of the Rockies specimen MOR 979, was collected from the Hell Creek of Montana, first published by American paleontologist John R. Horner , and preserves a nearly complete skull and skeleton. Boyd and colleagues also identified previously overlooked skull material of the T. neglectus paratype USNM 7758, which allowed comparisons of
484-477: 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
605-506: A larger clade of early ornithopods. In the description of T. assiniboiensis , identical results were recovered. Brown and colleagues found similar results within Ornithopoda again in 2013, prompting them to reintroduce the name Thescelosauridae for the total group, which could be divided into the revised subfamily Thescelosaurinae and the new subfamily Orodrominae . Thescelosaurus and Parksosaurus constituted Thescelosaurinae alongside
726-419: 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
847-412: A long, low snout that ended in a beak with a toothless tip. As in other dinosaurs, it was perforated by several fenestrae, or skull openings. Of these, the orbit (eye socket) and the infratemporal fenestra (behind the orbit) were proportionally large, while the external naris (nostril) was small. The external naris was formed by the premaxilla (the front bone of the upper jaw) and
968-489: A non-avian dinosaur, rivaling that of modern mammals such as the domestic cow . They reached their apex of diversity and ecological dominance in the hadrosaurids (colloquially known as 'duck-bills'), before they were wiped out by the Cretaceous–Paleogene extinction event along with all other non- avian dinosaurs . Members are known worldwide. In 1870, Thomas Henry Huxley listed Iguanodontidae (coined by Cope
1089-550: 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
1210-432: A primitive trait among ornithischians that is otherwise only found in much earlier and more basal forms such as Lesothosaurus and Scutellosaurus . Immature individuals may had less than six premaxillary teeth. Unlike many other basal ornithischians, the premaxillary teeth lacked serrations (small protuberances on the cutting edges). Both the maxilla and the dentary had up to twenty cheek teeth on each side, which
1331-608: 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
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#17327987201901452-515: A single specimen. Additional species have been suggested, but these are not widely considered valid, and some of them are only referable to the genus because the fossils are too incomplete to preserve the characteristic traits of any particular species. Thescelosaurus has been found across a wide geographic range. The type specimen was discovered in the Lance Formation of Wyoming , but subsequent discoveries of new specimens have expanded
1573-630: 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
1694-510: 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
1815-416: A year earlier ) as one of his three families of dinosaurs (alongside Megalosauridae and Scelidosauridae ), including within it the genera Iguanodon , Hypsilophodon , and Hadrosaurus , in addition to Cetiosaurus and tentatively Stenopelix . The term Ornithopoda was erected by Othniel Charles Marsh in 1881 as part of his then still ongoing investigation of the classification of Dinosauria. It
1936-570: 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
2057-418: 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
2178-427: Is again similar to basal ornithischians and unlike other neornithischians, which had a reduced tooth count. The cheek teeth themselves likewise showed primitive features, such as a constriction that separated the crowns from their roots , and a cingulum (bulge surrounding the tooth) above the constriction. The lower beak was formed by the predentary , a bone unique to ornithischians. When seen from below,
2299-656: Is also disagreement about whether Thescelosaurus and thescelosaurids are members of Ornithopoda, or more basal. Boyd highlighted in 2015 that many phylogenetic studies to include Thescelosaurus either do not include marginocephalians or are unresolved, so there was not definitive evidence that Thescelosaurus was an ornithopod. In his analysis, Thescelosaurus and Thescelosauridae were outside Ornithopoda, instead being an expansive clade of non-ornithopod neornithischians. Some studies agree with this placement for thescelosaurids, while others support Thescelosaurus as an ornithopod, and others are unresolved. Fonseca and colleagues gave
2420-633: Is best known from T. neglectus , mostly thanks to an almost complete example (specimen NCSM 15728) that has been CT-scanned to reveal its internal details. A fragmentary skull is also known from T. assiniboiensis (RSM P 1225.1). Most autapomorphies – distinguishing features that are not found in related genera – are found in the skull, as are most of the features that separate T. neglectus from T. assiniboiensis . The skull also shows many plesiomorphies , "primitive" features that are typically found in ornithischians which are geologically much older, but also shows derived (advanced) features. The skull had
2541-666: 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
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#17327987201902662-448: 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
2783-487: Is correct, and that if it is confirmed the species name garbanii should have priority. Isolated teeth from the Campanian Judith River Formation of Montana that were referred to Thescelosaurus cf. neglectus by Indian paleontologist Ashok Sahni in 1972, which would be the oldest occurrence of Thescelosaurus , were reassigned to Orodromeus in the same 1995 review. In a 1999 study on
2904-414: Is due to the anatomy of Parksosaurus being misinterpreted, and the anatomy of Thescelosaurus showing some variation, with Boyd demonstrating that Parksosaurus and Thescelosaurus were very closely related if not each others closest relatives. The clades Thescelosauridae (or alternatively Parksosauridae ) and Thescelosaurinae have been found by numerous phylogenetic analyses, though not all agree. There
3025-495: Is no direct evidence that Thescelosaurus lived in burrows or utilized digging as a foraging strategy, but a closely-related animal, Oryctodromeus , was fossilized inside a burrow, so these behaviors are known to have existed among similar small ornithischians during the Late Cretaceous. Researchers have also used computed tomography to examine the internal structure of the skull, and the braincase of Thescelosaurus
3146-494: Is often listed as an infraorder within a suborder Ornithopoda, though Benton (2004) lists Ornithopoda as an infraorder and does not rank Iguanodontia. Traditionally, iguanodontians were grouped into the superfamily Iguanodontoidea and family Iguanodontidae . However, phylogenetic studies show that the traditional "iguanodontids" are a paraphyletic grade leading up to the hadrosaurs (duck-billed dinosaurs). Groups like Iguanodontoidea are sometimes still used as unranked clades in
3267-512: Is usually given the rank of Suborder, within the order Ornithischia. While ranked taxonomy has largely fallen out of favour among dinosaur paleontologists, some researchers have continued to employ such a classification, though sources have differed on what its rank should be. Benton (2004) placed it as an infraorder within the suborder Cerapoda (originally named as an unranked clade ), while others, such as Ibiricu et al. 2010, have retained it at its traditional ranking of suborder. Iguanodontia
3388-496: The nasal bone , while the maxilla (the tooth-bearing "cheek" bone) was excluded. Another fenestra, the antorbital fenestra , was in-between the external naris and the orbit and contained two smaller internal fenestrae. Long rod-like bones called palpebrals were present above the eyes, giving the animal heavy bony eyebrows. The palpebral was not aligned with the upper margin of the orbit as in some other ornithischians, but protruded across it. The frontal bones , which form
3509-404: The skull roof above the orbit, were widest at the level of the middle of the orbit and narrower at their rear ends – an autapomorphy of Thescelosaurus . There was a prominent ridge along the length of both maxillae; a similar ridge was also present on both dentaries (the tooth-bearing bone of the lower jaw). The ridges and position of the teeth, deeply internal to the outside surface of
3630-512: The Cretaceous–Paleogene extinction event around 66 million years ago. Adult Thescelosaurus would have measured roughly 3–4 metres (10–13 ft) long and probably weighed several hundred kilograms. The genus Thescelosaurus is the type genus and also the largest member of the eponymous Thescelosauridae , which includes similarly-sized bipedal herbivores from the Late Cretaceous of Asia and North America such as Orodromeus , Parksosaurus , and Haya . Thescelosaurus bore several of
3751-511: The Greek ornithos , ornis ("bird") and pous , podos ("feet"); this is in reference to members’ characteristic birdlike feet. They were also characterized as lacking in body armour, not developing a horny beak , having an elongated pubis (that eventually extended past the ilium ), and having a missing hole in the lower jaw (a Mandibular fenestra ). A variety of ornithopods, and related ornithischians , had thin cartilaginous plates along
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3872-708: 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
3993-746: The Hell Creek Formation around Limas, Montana , AMNH 5889 found by Brown in 1806 in Hell Creek, Montana , CMN 9493 and 9507 found in 1921 by Canadian paleontologist Levi Sternberg in the Frenchman Formation of Rocky Creek, Saskatchewan , and CMN 9143 and 9534 also from Saskatchewan. AMNH 5034 was found only 5 ft (1.5 m) below the Fort Union Formation , as the youngest locality from which dinosaurs were found. From this material, Galton could not support
4114-831: The Hypsilophodontidae is problematic. The group previously consisted of all non- iguanodontian bipedal ornithischians, but a phylogenetic reappraisal has shown such species to be paraphyletic . As such, the hypsilophodont family is currently represented only by Hypsilophodon . Later ornithopods became larger, but never rivalled the incredible size of the long-necked, long-tailed sauropods that they partially supplanted. The very largest, such as Shantungosaurus , were as heavy as medium-sized sauropods (up to 23 metric tons /25 short tons ), but never grew much beyond 15 metres (50 feet). Historically, most indeterminate ornithischian bipeds were lumped in as ornithopods. Most have since been reclassified. Ornithopoda
4235-528: The K-Pg extinction . The fossil, consisting of a leg, was found to have particularly well-preserved skin. Thescelosaurus was a heavily built bipedal animal. Overall, the skeletal anatomy of this genus is well documented, and restorations have been published in several papers, including skeletal restorations and models. The skeleton is known well enough that a detailed reconstruction of the hip and hindlimb muscles has been made. The animal's size has been estimated in
4356-827: The Late Jurassic Kimmeridge Clay Formation of Weymouth, England , roughly 70 million years older than Bugenasaura and from another continent. Galton questioned whether the origins of the tooth were correct, as it had possibly been mislabelled and was actually from the Lance Formation of Wyoming, but the tooth was first collected before the UCMP was actively in the Lance region. The lack of diagnostic features led British paleontologists Paul M. Barrett and Susannah Maidment to classify UCPM 49611 as an indeterminate ornithischian in 2011. With
4477-567: The Royal Ontario Museum as ROM 804, was found around 0.5 mi (0.80 km) east of the Red Deer River , 100 ft (30 m) above the water level. The similarity to Thescelosaurus prompted Canadian paleontologist William A. Parks to describe ROM 804 in 1926 as the new species Thescelosaurus warreni , named after ROM Board of Trustees member H.D. Warren, as the first member of Hypsilophodontidae from Canada and
4598-501: 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)
4719-488: The skull roof and a complete mandible , the first known from Thescelosaurus . Sternberg also reconsidered the separation of Thescelosauridae, instead proposing the subfamily Thescelosaurinae to house Thescelosaurus separately from Parksosaurus , Hypsilophodon and Dysalotosaurus within Hypsilophodontidae. Thescelosaurus was reviewed by American paleontologist Peter M. Galton in 1974, including
4840-506: The theropods , to help them balance as they ran on their hind legs. Later ornithopods became more adapted to grazing on all fours; their spines curved, and came to resemble the spines of modern ground-feeders, such as the bison . As they became more adapted to eating while bent over, they became facultative quadrupeds; still running on two legs, and comfortable reaching up into trees, but spending most of their time walking or grazing on all fours. The taxonomy of dinosaurs previously ascribed to
4961-410: The 2.5–4.0 m range for length (8.2–13.1 ft) for various specimens, and a weight of 200–300 kilograms (450–660 pounds ), with the large type specimen of T. garbanii estimated at 4–4.5 meters (13.1–14.8 feet) long. It may have been sexually dimorphic , with one sex larger than the other. Juvenile remains are known from several locations, mostly based on teeth. The skull
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5082-1031: 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"):
5203-495: 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
5324-514: The Asian taxa Haya griva , Jeholosaurus and Changchunsaurus that had been found to be related previously, with Zephyrosaurus , Orodromeus , Oryctodromeus , the new genus Albertadromeus , and an unnamed specimen forming Orodrominae. Some studies separate from Boyd and Brown did not find Parksosaurus to be closely related to Thescelosaurus , instead forming a relationship with South American Gasparinisaura . However, this relationship
5445-654: 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, Ornithopoda Ornithopoda ( / ˌ ɔːr n ə ˈ θ ɒ p ə d ə / ) is a clade of ornithischian dinosaurs , called ornithopods ( / ˈ ɔːr n ə θ ə ˌ p ɒ d z , ɔːr ˈ n ɪ θ -/ ). They represent one of
5566-518: The Cretaceous, but these animals were generally smaller and less robust than Thescelosaurus itself. The genus attracted media attention in 2000, when a specimen unearthed in 1993 in South Dakota , United States, was interpreted as including a fossilized heart. There was much discussion over whether the remains were of a heart. Many scientists now doubt the identification of the object and
5687-645: The European Hypsilophodon and three American taxa he named himself, Camptonotus , Laosaurus , and Nanosaurus . Camptonotus was in 1885 renamed to Camptosaurus , as the original name was pre-occupied by a cricket ; the associated family followed suit, becoming Camptosauridae. In Iguanodontidae, only found in Europe, he included Iguanodon and Vectisaurus . In Hadrosauridae, he included Hadrosaurus , Cionodon , and tentatively Agathaumas . Ornithopoda means "bird feet", from
5808-682: The Hell Creek Formation of Garfield County, Montana by Harli Garbani and stored in the Los Angeles County Museum of Natural History , and one found in an unknown location within Harding County, South Dakota and stored in the South Dakota School of Mines and Technology Geology Museum. The first specimen, LACM 33543, preserved parts of the vertebral column and pelvis in addition to bones of
5929-719: The Lance and Hell Creek Formations, T. assiniboiensis from the Frenchman Formation, T. garbanii from the Hell Creek Formation, and other indeterminate specimens across all localities. In April 2022, news media reported that a specimen of Thescelosaurus was found at the Tanis fossil site in North Dakota. This site is thought to show direct signs of the Chicxulub asteroid impact in the Gulf of Mexico that resulted in
6050-565: The Tanis specimen preserves skin impressions on a leg that show that parts of the animal were covered in scales. While Thescelosaurus was originally considered a member of Camptosauridae by Gilmore alongside Hypsilophodon , Dryosaurus and Laosaurus , he revised his opinion following further study to place the taxon within Hypsilophontidae (a misspelling of Hypsilophodontidae) alongside only Hypsilophodon . Many authors followed
6171-882: The Tolman Member of the Horseshoe Canyon Formation between 70.896 and 69.6 million years old, and Thescelosaurus from the Scollard Formation between 66.88 million years old and the end of the Cretacous. Sternberg described the Thescelosaurus specimen, accessioned in the Canadian Museum of Nature as CMN 8537, in 1940 as a new species, T. edmontonensis , known from most of the vertebral column , pelvis, legs, scapula, coracoid, arm, and most significantly multiple bones of
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#17327987201906292-412: The ability to chew, and Thescelosaurus conforms to this pattern. Analysis of their teeth suggests that they may have been more selective in feeding than similarly-sized pachycephalosaurids . Thescelosaurus also has a number of adaptations which have been interpreted as signs of fossoriality . These include robust arms, large olfactory bulbs, reduced hearing capacity, and shortened hindlimbs. There
6413-414: The anatomy of Bugenasaura , Galton referred two isolated teeth to the genus, one ( Yale Peabody Museum 8098) collected by Hatcher in 1889 from the Lance Formation of Lusk, Wyoming , was referred to B. infernalis , while another, University of California Museum of Paleontology 49611 was referred to cf. Bugenasaura , as it showed some anatomical differences, but most significantly was listed as being from
6534-491: The animal aged. Such plates are known from several other cerapodans . The front ribs were flattened and concave, and the rear margins of their lower ends had a rough surface. These features are autapomorphies of Thescelosaurus and are possibly adaptations that allow the plates to attach to the rib cage. For most of its history, the nature of this genus' integument , be it scales or something else, remained unknown. Charles Gilmore described patches of carbonized material near
6655-550: 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
6776-419: 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
6897-435: The body, another idea that has gone by the wayside. As noted by Peter Galton , the upper arm bone of most ornithischians articulated with the shoulder by an articular surface that consisted of the entire end of the bone, instead of a distinct ball and socket as in mammals. The orientation of the shoulder's articular surface also indicates a vertical and not horizontal upper arm in dinosaurs. The species T. garbanii differs from
7018-438: 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
7139-641: The characteristic traits of ornithischian dinosaurs including a beak, an inverted pubis , a relatively small antorbital fenestra , a closed mandibular fenestra , and five digits on the hands. However, the animals bore a mix of derived and basal traits when compared with similarly-sized ornithischians. The tip of the snout was edentulous , but the premaxillary , maxillary , and dentary bones bore both pointed and leaf-shaped serrated teeth. It also possessed tridactyl feet similar to those of theropods and ornithopods . The tail of Thescelosaurus bore rod-like ossified tendons , which aided in counterbalancing
7260-452: The classification of Thescelosaurus within Hypsilophodontidae, but not all. Hungarian paleontologist Franz Nopcsa and German paleontologist Friedrich von Huene retained Thescelosaurus (misspelled "Thescelesaurus" by Nopcsa) as a relative of Camptosaurus . Sternberg at first separated Thescelosaurus and related Parksosaurus into Thescelosauridae, before considering both members of Hypsilophodontidae with Thescelosaurus separated from
7381-513: The diagnostic regions of the skull and ankle across multiple specimens and species. Based on the limitations of overlapping material, T. neglectus was restricted to only the types USNM 7757 and 7758, and T. garbanii was maintained as a species of Thescelosaurus but limited to its type LACM 33542, all other specimens, including the types of T. edmontonensis and Bugenasaura infernalis , were referred to Thescelosaurus incertae sedis , as they showed features characteristic of Thescelosaurus in
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#17327987201907502-615: 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
7623-585: The discovery of additional specimens of Thescelosaurus preserving both the skull and skeleton, American paleontologist Clint Boyd and colleagues reassessed the historic and current species of Thescelosaurus in 2009. One of the new specimens, housed in the North Carolina State Museum of Natural Sciences (NCSM 15728), was found in the upper Hell Creek Formation in Harding County, South Dakota by Michael Hammer in 1999, and preserves
7744-569: The first description of the material in the AMNH mentioned by Gilmore in 1915. At the time of Galton's review, 15 specimens of Thescelosaurus were available to be described, and additional material was being worked on by American paleontologist William J. Morris. In addition to the specimens previously described, there was AMNH 117 found in 1892 by Wortman and Peterson at an uncertain location, AMNH 5031, 5034 and 5052 found by American paleontologists Barnum Brown and Peter Kaisen in 1909 from localities of
7865-548: The first specimen of Thescelosaurus to preserve a skull. Also in 1926, Canadian paleontologist Charles Mortram Sternberg noted the discovery of a new specimen of Thescelosaurus , also from the Edmonton Formation, 7 mi (11 km) northwest of Rumsey, Alberta and 260 ft (79 m) above the water level of the Red Deer River. The specimen, found within the last three field seasons by Sternberg,
7986-673: 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
8107-561: 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
8228-476: 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
8349-430: The identification of T. edmontonensis as a separate species, instead considering all the specimens to represent T. neglectus . The few differences identified between T. edmontonensis and all the other material were considered to represent either individual or sexual variation, not significant enough to justify a separate species. Morris published his description of three Thescelosaurus specimens in 1976, two found in
8470-531: The implications of such an identification. In July of 1891 American paleontologists John Bell Hatcher and William H. Utterback discovered a partial skeleton of a small ornithopod in Wyoming . The specimen was found in Doegie Creek, Niobrara County , which was at the time part of Converse County , and was nearly complete and articulated, lacking only the skull and neck, and parts of the arm bones. It
8591-458: The most primitive hypsilophodontid outside the remaining taxa. The analysis of Weishampel and Heinrich in 1992 can be seen below. Thescelosaurus Yandusaurus Othnielia Parksosaurus Hypsilophodon Zephyrosaurus Orodromeus The concept of Hypsilophodontidae as a monophyletic group then fell out of favor, instead it was suggested in 1999 by American paleontologist Rodney Sheetz that "hypsilophodontids" were simply
8712-466: The most successful groups of herbivorous dinosaurs during the Cretaceous . The most primitive members of the group were bipedal and relatively small-sized, while advanced members of the subgroup Iguanodontia became quadrupedal and developed large body size. Their major evolutionary advantage was the progressive development of a chewing apparatus that became the most sophisticated ever developed by
8833-714: The name Pyrodontia to the clade uniting Thescelosaurus with more derived ornithischians when Thescelosauridae is outside Ornithopoda, referencing the early and rapid diversification of Thescelosauridae, Marginocephalia and Ornithopoda. The thescelosaurid results of Fonseca and colleagues in 2024 can be seen below. Zephyrosaurus Oryctodromeus Orodromeus Koreanosaurus Albertadromeus Yueosaurus Changmiania Haya griva RTMP 2008.045.0002 Jeholosaurus Changchunsaurus Parksosaurus CMN 8537 ( T. edmontonensis ) T. neglectus Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
8954-435: The neck and back, and a nearly complete lower leg with a partial femur . Morris identified that the larger size and ankle of this specimen was unique, with the reduction of the calcaneum bone, and as such he named it Thescelosaurus garbanii , in honor of the discoverer and preparator Garbani. Morris also suggested that the ankle of T. edmontonensis had been previously misinterpreted and was more similar to T. garbanii in
9075-545: The neck and skull to illustrate the complete skeleton of Thescelosaurus neglectus . The type skeleton of Thescelosaurus was first displayed in the Hall of Extinct Monsters of the Smithsonian National Museum of Natural History (formerly United States National Museum), but was then redisplayed in 1963 more prominently as a wall-mount alongside the ornithischians Edmontosaurus and Corythosaurus and
9196-600: The neck vertebra USNM 7761 found in 1891 by Hatcher, Sullins and Burrell in Beecher's Quarry in Niobrara County, the phalanx of the foot USNM 8065 found in 1890 by Hatcher in Niobrara County, and three undescribed partial skeletons at the AMNH found in Dawson County, Montana . Thescelosaurus was found to be more similar to Hypsilophodon than Camptosaurus by Gilmore in 1915, from which he reconstructed
9317-410: The neck, back, and tail, parts of the pectoral girdle and arm, and a partial leg. It was collected in 1889 by Olof August Peterson at Lance Creek , Niobrara County. Both Lance Creek and Doegie Creek localities are part of the Lance Formation , which is a Maastrichtian deposit that spans from 69.42 million years ago until the end of the Cretaceous . Preparation of the type specimen of Thescelosaurus
9438-462: The new taxon Bugenasaura infernalis . The name was a combination of the Latin bu , "large", gena , "cheek", and saura , "lizard", with the species name as a reference to the lower levels of the Hell Creek Formation from which it is known. Galton also tentatively referred LACM 33543, the type of T. garbanii , to the new species, noting that additional material is necessary to determine if the referral
9559-508: 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
9680-429: The ornithopod family Camptosauridae , so in 1913 he published a preliminary description naming the specimen Thescelosaurus neglectus ; the genus name derived from the Greek words θέσκελος ( theskelos ), "marvelous", and σαυρος ( sauros ) "reptile" or "lizard". As well as the type specimen USNM 7757, Gilmore referred USNM 7758 to Thescelosaurus in 1913, a partial skeleton including vertebrae from
9801-418: The other genera as a member of Thescelosaurinae. Russian paleontologist Anatoly Konstantinovich Rozhdestvensky and Australian paleontologist Richard A. Thulborn retained Thescelosauridae as a separate family. Galton argued against the inclusion of Thescelosaurus within Hypsilophodontidae originally, instead emphasizing the hindlimb proportions in classifying it as a member of Iguanodontidae . Iguanodontidae
9922-400: The other species in its unique ankle, with the calcaneus being reduced and not contributing to the midtarsal joint . T. neglectus had 6 sacral ("hip") vertebrae and 27 presacral ("neck and back") vertebrae. The holotype specimen of the species T. assiniboiensis appears to have had only five sacrals, but it is possible that this individual was not yet fully mature and that the last sacral
10043-415: The outside of the ribs; in some cases, these plates mineralized and were fossilized. The function of these intercostal plates is unknown. They have been found with Hypsilophodon , Nanosaurus , Parksosaurus , Talenkauen , Thescelosaurus , and Macrogryphosaurus to date. The early ornithopods were only about 1 metre (3 feet) long, but probably very fast. They had a stiff tail, like
10164-537: The paraphyletic nature of Hypsilophodontidae . A 2017 study which named and described Burianosaurus noted that the type species Iguanodon bernissartensis must be part of the definition, and that the 2005 definition would, in their analysis, include a far larger group than intended (including Marginocephalia ). They proposed an entirely new, node-based definition: the last common ancestor of Iguanodon bernissartensis , Dryosaurus altus , Rhabdodon priscus , and Tenontosaurus tilletti . In 2021, Iguanodontia
10285-487: 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
10406-419: The primitive form of ornithopods. Scheetz found Thescelosaurus , Parkosaurus and Bugenasaura to be successively closer to Hypsilophodon and later ornithopods, but not a group of their own. Such a result became common, with Thescelosaurus or Bugenasaura as an early ornithopod close to the origins of the group, sometimes forming a clade with Parksosaurus . An issue with Thescelosaurus neglectus prior to
10527-544: The range of Thescelosaurus to include North Dakota , South Dakota , Montana , Alberta , and Saskatchewan . All of the places where Thescelosaurus remains have been found — the Frenchman Formation , Hell Creek Formation , Scollard Formation , and others — have been dated to the later part of the Maastrichtian (the very end of the Cretaceous Period). Relatives of Thescelosaurus lived earlier in
10648-405: The rear end of the predentary was bifurcated, which is a derived feature. Thescelosaurs had short, broad, five-fingered hands, four-toed feet with hoof -like toe tips , and a long tail braced by ossified tendons from the middle to the tip, which would have reduced the flexibility of the tail. The rib cage was broad, giving it a wide back, and the limbs were robust. The femur (upper thigh bone)
10769-417: The reduction of the calcaneum, and that the two species may eventually be separated from Thescelosaurus as a new genus. The third specimen, SDSM 7210, was provisionally referred to Thescelosaurus as it includes a large part of the skull, some partial vertebrae from the back and two bones of the fingers, preventing comparison with the diagnostic regions of the Thescelosaurus species. Morris chose not to name
10890-449: The review of Boyd and colleagues in 2009 was the uncertainty about the referred materials, including the separation of Bugenasaura and lack of clear synonymy of T. edmontonensis . Following the taxonomic revision, the systematic relationships of Thescelosaurus and "hypsilophodonts" have become clearer, with Boyd and colleagues finding Thescelosaurus and Parksosaurus to unite with Zephyrosaurus , Orodromeus and Oryctodromeus as
11011-470: The scientific literature, though many traditional "iguanodontids" are now included in the more inclusive group Hadrosauroidea . Iguanodontia was originally phylogenetically defined, by Paul Sereno , in 1998, as the most inclusive group containing Parasaurolophus walkeri but not Hypsilophodon foxii . Later, in 2005, he amended the definition to include Thescelosaurus neglectus as a secondary external specifier, alongside Hypsilophodon , accounting for
11132-439: The shoulders as possible epidermis , with a "punctured" texture, but no regular pattern, while William J. Morris suggested that armor was present, in the form of small scutes he interpreted as located at least along the midline of the neck of one specimen. Scutes have not been found with other articulated specimens of Thescelosaurus , though, and Morris's scutes could be crocodilian in origin. In 2022, news media reported that
11253-483: The skull and hindlimb, but either did not preserve the squamosal bone of the skull for comparison to T. neglectus , or the ankle for comparison to T. garbanii . With the referral of Bugenasaura to Thescelosaurus , Boyd and colleagues created the new combination T. infernalis , but could not identify features of the skull to distinguish the species from others, considering it undiagnostic. Two specimens were noted as having anatomy slightly different from T. neglectus ,
11374-480: The skull now yet known from Thescelosaurus such as the jugals and braincase . Morris referred this specimen to T. neglectus , and also noted that two small scutes were found above the neck vertebrae of this individual. These scutes were suggested to be crocodilian and not from Thescelosaurus by Galton in 2008, and no other specimens that preserve the neck suggest osteoderms were present. The second specimen described by Morris, LACM 33542, includes vertebrae from
11495-415: The skull, are interpreted as evidence for muscular cheeks. The morphology of the ridge on the maxilla, which is very pronounced and has small and oblique ridges covering its rear end, is an autapomorphy of the genus. The teeth were of two types : small pointed premaxillary teeth (in the premaxilla, or upper beak), and leaf-shaped cheek teeth (in the maxilla and the dentary). The premaxillae had six teeth each,
11616-458: The specimen due to this lack of overlapping material, but regarded the specimen as a new species nonetheless. American paleontologist Hans-Dieter Sues agreed with retaining SDSM 7210 as unnamed in his description of the hypsilophodontid Zephyrosaurus , informally referring to it as the Hell Creek hypsilophodontid. The species of Thescelosaurus were reviewed again by Galton in 1995. The diagnostic ankle of T. edmontonensis identified by Morris
11737-454: The specimens NCSM 15728 and Royal Saskatchewan Museum specimen RSM P 1225.1 (previously described by Galton in 1995 and 1997 as T. neglectus ). However, the ankle of NCSM 15728 is unknown, preventing separation from T. garbanii , and the skull of RSM P 1225.1 showed some similarity to T. edmontonensis , which would be the proper name for the species if further study showed both specimens could be distinguished from T. neglectus . Parksosaurus
11858-517: 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
11979-467: The theropod Gorgosaurus , a collection of dinosaurs that did not live at the same time or in the same place. In 1981 the display was rearranged placing Thescelosaurus higher and more out-of-sight. Renovations of the exhibit from 2014 to 2019 removed the Thescelosaurus and other dinosaurs on display, replacing them with remounted casts so that the original fossils could be further prepared and studied. The duplication and repreparation of Thescelosaurus
12100-438: The weight of the animal. The ribs of Thescelosaurus were accompanied by thin plate-like mineralized tissues, the precise function of which is not confidently known. Thescelosaurus is relatively unique among thescelosaurids for having a relatively long femur in relation to the tibia , suggesting that it was probably not a very fast runner. All known ornithischians are presumed to have been primarily herbivorous and possessed
12221-480: 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 ,
12342-432: Was also the possibility that the hindlimb of T. garbanii was instead not a hypsilophodontid, but the hindlimb of the pachycephalosaurid Stygimoloch also known from the Hell Creek Formation, for which the hindlimb was unknown. Following the reassessments of T. edmontonensis and T. garbanii , Galton concluded that the skull of SDSM 7210 was distinct from all other hypsilophodontids including Thescelosaurus , and named
12463-652: Was completed between 1913 and 1915, at which point Gilmore published a full monograph of the taxon, and identified six more specimens in the collections of the USNM and the American Museum of Natural History that could be identified as the same species. These additional specimens include the scapula and coracoid USNM 7760 found in 1891 by Hatcher in Deer Ears Buttes in Butte County, South Dakota ,
12584-508: Was considered one of the four definite orders of dinosaurs, the others being Theropoda , Sauropoda , and Stegosauria ( Hallopoda was considered a possible fifth). He subdivided the order into three families: Camptonotidae , Iguanodontidae , and Hadrosauridae ; the former was a new name, whereas the latter two were carried over from the nomenclatures of Huxley and Edward Drinker Cope respectively. Within Camptonotidae he included
12705-575: Was determined that it represented a new species they named T. assiniboiensis . The species name derives from the historic District of Assiniboia that covered the southern Saskatchewan region where the Frenchman Formation is exposed, which in turn takes its name from the Assiniboine peoples. The presence of a foramen in the braincase not found in any other specimens of Thescelosaurus , including CMN 8537 (type of T. edmontonensis ) and NCSM 15728. The separation of T. assiniboiensis and T. neglectus
12826-543: Was done by Research Casting International, who replicated the ossified tendons and cartilage of the original fossil in the mount, and updated the skull to a Thescelosaurus specimen described in 2014. A skeleton of a similar ornithopod to Thescelosaurus neglectus was found in 1922 by an expedition of the University of Toronto to the Edmonton Formation of Alberta . This specimen, collected and taken to
12947-530: Was expanded by American paleontologist Alfred Sherwood Romer to include most small ornithopods by 1966, which was followed by Galton (though Thescelosaurus was removed) and later authors, with Hypsilophodontidae including 13 genera in the first edition of the book The Dinosauria in 1990. This concept of Hypsilophodontidae as an expansive natural group has been recovered by the early cladistic studies of American paleontologists Paul C. Sereno , and David B. Weishampel and Ronald Heinrich, with Thescelosaurus as
13068-484: Was from the northwest side of a butte on the north side of the valley, approximately halfway up the exposed claystone . This places the specimen in the Frenchman Formation, which was deposited in the last half million years before the end of the Cretaceous. The specimen, described as T. neglectus by Galton, and an indeterminate species requiring further study by Boyd and colleagues, was fully studied by Canadian paleontologist Caleb M. Brown and colleagues in 2011, where it
13189-523: Was further supported by Boyd in his 2014 description of the skull of NCSM 15728 and Timber Lake and Area Museum specimen TLAM.BA.2014.027.0001, discovered and collected from private lands by Bill Alley before being donated to the museum, which had yet to be fully prepared but includes a mostly complete but slightly crushed skull and much of the skeleton. Thescelosaurus is known from multiple specimens found throughout Alberta, Saskatchewan, Wyoming, North Dakota, South Dakota and Montana, with T. neglectus from
13310-590: Was given a formal definition under the PhyloCode : "The smallest clade containing Dryosaurus altus , Iguanodon bernissartensis , Rhabdodon priscus , and Tenontosaurus tilletti , provided that it does not include Hypsilophodon foxii ." Under this revised definition, Iguanodontia is limited to its traditionally included species, and if it were found to include hypsilophodonts, which were not traditionally considered iguanodontians, it would become an invalid grouping. The slightly less inclusive clade Dryomorpha
13431-507: Was identified as a " hypsilophodont " — a wastebasket taxon that included a variety of unrelated, but superficially similar bipedal ornithischians . Modern taxonomists consider Thescelosaurus to be either a stem-neornithischian, or a basal member of Ornithopoda , although its precise affinities remain somewhat uncertain. The genus also contains at least two other valid species: T. garbanii and T. assiniboiensis , which were named in 1976 and 2011, respectively and are each known from
13552-441: Was likely complete when buried. This posture was maintained for the exhibit of the skeleton, with only the right leg, which was slightly dislocated, being adjusted in position. Some minor damage to the bones was restored, but painted lighter than the original bones so that the real and reconstructed parts could be distinguished visually. From the legs, pelvis and hands of the skeleton, Gilmore found it unique from all other members of
13673-418: Was longer than the tibia (shin bone), which distinguishes the genus from closely related genera. The animals may have been able to move on all fours , given its fairly long arms and wide hands, but this idea has not been widely discussed in the scientific literature, although it does appear in popular works. Charles M. Sternberg reconstructed it with the upper arm held horizontally and almost perpendicular to
13794-613: Was named by Paul Sereno in 1986 and given a formal definition in the PhyloCode as "the smallest clade containing Dryosaurus altus and Iguanodon bernissartensis ". This group includes basal members such as Hesperonyx , members of the family Dryosauridae , and the derived clade Ankylopollexia . In 2021, Ornithopoda was given a formal definition under the PhyloCode : "The largest clade containing Iguanodon bernissartensis but not Pachycephalosaurus wyomingensis and Triceratops horridus ." The cladogram below follows
13915-486: Was not considered to have evolved from a single evolutionary source, but instead composed a polyphyletic group of ornithopods with similar convergent bauplans. However, he revised his taxonomy of Thescelosaurus in 1995, returning to a hypsilophodontid classification. Hypsilophodontidae only included four genera when its classification was assessed by Sternberg in 1940, Hypsilophodon , Thescelosaurus , Parksosaurus , and Dysalotosaurus . The content of Hypsilophodontidae
14036-466: Was not yet fused to the other sacrals. Large thin flat mineralized plates have been found next to the ribs' sides. Their function is unknown; they may have played a role in respiration . However, muscle scars or other indications of attachment have not been found for the plates, which argues against a respiratory function. Recent histological study of layered plates from a probable subadult indicates that they may have started as cartilage and became bone as
14157-495: Was not yet prepared, but did include parts of the skull. As there were substantial differences between T. warreni , T. neglectus and the other Edmonton specimen, Sternberg placed T. warreni in the new genus Parksosaurus in 1937, and proposed the new family Thescelosauridae to unite the two genera. Newer geology has separated the Edmonton Formation into four formations as the Edmonton Group , with Parksosaurus from
14278-482: Was reinterpreted as the result of breakage on the right ankle, with the previously undescribed left ankle showing the same anatomy as T. neglectus . As a result, Galton synonymized T. edmontonensis with T. neglectus again. Galton determined that Morris correctly interpreted the ankle of T. garbanii , preventing it from being a synonym of T. neglectus , but suggesting it is different enough to be an entirely separate genus of hypsilophodontid from Thescelosaurus . There
14399-493: Was relatively small for its size, suggesting that the animal was more comparable to squamates and crocodilians in intelligence than to other ornithischians. The type species of Thescelosaurus , T. neglectus , was described in 1913 by scientists working for the Smithsonian Institution , and numerous specimens have been referred to the type species in the century since it was described. Initially, it
14520-628: Was retained as a separate genus. RSM P 1225.1 was first found on June 19th, 1968 by Albert Swanson of the Saskatchewan Museum of Natural History (now the Royal Saskatchewan Museum), who collected the specimen on July 17th. The originally reported location was incorrect, revisiting of the Frenchman River valley by Tim Tokaryk in the 1980s found that the excavation, identifiable by bone and plaster remnants,
14641-509: Was taken to the United States National Museum and accessioned as specimen USNM 7757, where it remained in the original packing boxes until close to 1913 when it was identified as a new taxon by American paleontologist Charles W. Gilmore . When the skeleton was found, it was lying on its left side with nearly all bones articulated, but the skull, neck, and parts of the pectoral girdle had been eroded away, though it
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