112-516: Pachycephalosauria ( / ˌ p æ k ɪ s ɛ f əl ə ˈ s ɔː r i ə , - ˌ k ɛ f -/ ; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs . Along with Ceratopsia , it makes up the clade Marginocephalia . With the exception of two species, most pachycephalosaurs lived during the Late Cretaceous Period, dating between about 85.8 and 66 million years ago. They are exclusive to
224-543: 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
336-463: A 2009 petition to the ICZN . Majungasaurus was a medium-sized theropod that typically reached 5.6–7 m (18–23 ft) in length and weighed 750–1,100 kg (1,650–2,430 lb). Fragmentary remains of larger individuals indicate that some adults could have been similar in size to its relative Carnotaurus , possibly exceeding 8 m (26 ft) in length. The skull of Majungasaurus
448-568: A basal pachycephalosaur. The oldest known definitive pachycephalosaur is Sinocephale bexelli from the Late Cretaceous of China . In 2017, a phylogenetic analysis conducted by Han and colleagues identified Stenopelix as a member of the Ceratopsia . Pachycephalosaurs were bipedal ornithischians characterized by their thickened skulls. They had a bulky torso with an expanded gut cavity and broad hips, short forelimbs, long legs,
560-472: A close relationship with Carnotaurus from South America, while others were unable to firmly place it in the phylogeny . The most recent analysis, using the most complete information, instead recovered Majungasaurus in a clade with Rajasaurus and Indosaurus from India, but excluding South American genera like Carnotaurus , Ilokelesia , Ekrixinatosaurus , Aucasaurus and Abelisaurus , as well as Rugops from mainland Africa. This leaves open
672-399: A dome-shaped swelling nearly identical to the one described by Sues and Taquet as Majungatholus atopus . Majungatholus was redescribed as an abelisaurid rather than a pachycephalosaur in 1998. Although the name Majungasaurus crenatissimus was older than Majungatholus atopus , the authors judged the type dentary of Majungasaurus too fragmentary to confidently assign to the same species as
784-414: A group with Stegoceras as one of the earliest fully-domed members, with flat-headed and potentially juvenile taxa like Homalocephale and Goyocephale either just outside or just within the clade of it and more derived pachycephalosaurs. These studies began with the phylogenetic work of Paul Sereno , which has been modified in many iterations to include newer taxa and additional characters. The version of
896-407: A high degree of flexibility in the lower jaw, although not to the extent seen in snakes . This may have been an adaptation to prevent the fracture of the lower jaw when holding onto a struggling prey animal. The front teeth of the upper jaw were more robust than the rest, to provide an anchor point for the bite, while the low crown height of Majungasaurus teeth prevented them from breaking off during
1008-477: 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
1120-419: 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
1232-487: A pachycephalosaurid, has also since been reclassified as a coelurosaur (Naish in Sullivan (2006) ). Further complicating matters are the diverse interpretations of ontogenetic and sexual features in pachycephalosaurs. A 2009 paper proposed that Dracorex and Stygimoloch were just early growth stages of Pachycephalosaurus , rather than distinct genera. A 2020 reworking of Cerapoda by Dieudonné et al. recovered
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#17327903659661344-550: 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
1456-657: A re-evaluation of pachycephalosaur taxonomy. Sullivan considered attempts by Maryańska and Osmólska to restrict the definition of Pachycephalosauria redundant with their Pachycephalosauridae, since they were diagnosed by the same anatomical characters. Sullivan also rejected attempts by Sereno (1986) , in his phylogenetic studies, to re-define Pachycephalosauridae to include only "dome-skulled" species (including Stegoceras and Pachycephalosaurus ), while leaving more "basal" species outside that family in Pachycephalosauria. Therefore, Sullivan's use of Pachycephalosauridae
1568-637: A result of the harsh environment in which it lived. However, other abelisaurids have also been found to have comparably slow growth rates. All specimens of Majungasaurus have been recovered from the Maevarano Formation in the Mahajanga Province in northwestern Madagascar. Most of these, including all of the most complete material, came from the Anembalemba Member, although Majungasaurus teeth have also been found in
1680-608: 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
1792-538: A short, thick neck, and a heavy tail. Large orbits and a large optic nerve point to pachycephalosaurs having good vision, and uncharacteristically large olfactory lobes indicate that they had a good sense of smell relative to other dinosaurs. They were fairly small dinosaurs, with most falling in the range of 2–3 meters (6.6–9.8 feet) in length and the largest, Pachycephalosaurus wyomingensis , estimated to measure 4.5 meters (14.8 feet) long and weigh 450 kilograms (990 pounds). The characteristic skull of pachycephalosaurs
1904-421: A small flocculus, did not rely on quick head movements to sight and capture prey. Inferences about behavior can also be drawn from examination of the inner ear. The semicircular canals within the inner ear aid in balance, and the lateral semicircular canal is usually parallel to the ground when the animal holds its head in an alert posture. When the skull of Majungasaurus is rotated so that its lateral canal
2016-420: A solid bony mass. There is no sign of any other vertebrae after the fifth in the series, indicating that the tail ended there prematurely. From the size of the last vertebrae, scientists judged that about ten vertebrae were lost. One explanation for this pathology is severe physical trauma resulting in the loss of the tail tip, followed by osteomyelitis (infection) of the last remaining vertebrae. Alternatively,
2128-630: 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
2240-402: A struggle. Finally, unlike the teeth of Allosaurus and most other theropods, which were curved on both the front and back, abelisaurids like Majungasaurus had teeth curved on the front edge but straighter on the back (cutting) edge. This structure may have served to prevent slicing, and instead holding the teeth in place when biting. Examination of the teeth of Majungasaurus indicates that
2352-513: A very strong and muscular neck . Uniquely, the cervical ribs of Majungasaurus had long depressions along the sides for weight reduction. The humerus (upper arm bone) was short and curved, closely resembling those of Aucasaurus and Carnotaurus . Also like related dinosaurs, Majungasaurus had very short forelimbs with four extremely reduced digits, first reported with only two very short external fingers and no claws. The hand and finger bones of Majungasaurus , like other majungasaurines, lacked
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#17327903659662464-510: 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
2576-594: Is a genus of abelisaurid theropod dinosaur that lived in Madagascar from 70 to 66 million years ago , at the end of the Cretaceous Period , making it one of the last-known non-avian dinosaurs that went extinct during the Cretaceous–Paleogene extinction event . The genus contains a single species , Majungasaurus crenatissimus . This dinosaur is also called Majungatholus , a name which
2688-858: Is a cladogram published by Dieudonné and colleagues (2020) which controversially found heterodontosauridae to be paraphyletic with respect to pachycephalosauria. This analysis was proposed as a hypothesis for the complete lack of Jurassic and Early Cretaceous pachycephalosaur fossils, even though they should have existed if the modern understanding of ornithischian phylogeny is correct. However, this hypothesis has not been widely accepted by other paleontologists. Fruitadens Lycorhinus Heterodontosaurus Abrictosaurus Tianyulong Echinodon Wannanosaurus Goyocephale Prenocephale Homalocephale Stegoceras Pachycephalosaurus The small size of most pachycephalosaur species and lack of skeletal adaptation indicates that they were not climbers and primarily ate food close to
2800-570: 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
2912-470: Is a result of the fusion and thickening of the frontals and parietals , accompanied by the closing of the supratemporal fenestra . In some species this takes the form of a raised dome; in others, the skull is flat or wedge-shaped. While the flat-headed pachycephalosaurs are traditionally regarded as distinct species or even families, they may represent juveniles of dome-headed adults. All display highly ornamented jugals , squamosals , and postorbitals in
3024-418: 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
3136-666: 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
3248-431: Is considered a junior synonym of Majungasaurus . Like other abelisaurids, Majungasaurus was a bipedal predator with a short snout. Although the forelimbs are not completely known, they were very short, while the hind limbs were longer and very stocky. Measuring around 7 m (23 ft) long and weighing more than 1,000 kg (2,200 lb), it can be distinguished from other abelisaurids by its wider skull ,
3360-448: 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
3472-578: Is direct evidence of cannibalism . French paleontologist Charles Depéret described the first theropod remains from northwestern Madagascar in 1896. These included two teeth, a claw, and some vertebrae discovered along the Betsiboka River by a French army officer and deposited in the collection of what is now the Université Claude Bernard Lyon 1 . Depéret referred these fossils to the genus Megalosaurus , which at
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3584-671: Is equivalent to Sereno and Benton's use of Pachycephalosauria. Sullivan diagnosed the Pachycephalosauridae-based only on characters of the skull, with the defining character being a dome-shaped frontoparietal skull bone. According to Sullivan, the absence of this feature in some species assumed to be primitive led to the split in classification between domed and non-domed pachycephalosaurs; however, discovery of more advanced and possibly juvenile pachycephalosaurs with flat skulls (such as Dracorex hogwartsia ) show this distinction to be incorrect. Sullivan also pointed out that
3696-408: Is evidence that the dome had some form of external covering, and it is reasonable to consider the dome may have been brightly covered, or subject to change color seasonally. Due to the nature of the fossil record, however, it cannot be observed whether or not color played a role in dome function. Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
3808-467: Is exceptionally well known compared to most theropods and generally similar to that of other abelisaurids. Like other abelisaurid skulls, its length was proportionally short for its height, although not as short as in Carnotaurus . The skulls of large individuals measured 60–70 centimeters (24–28 in) long. The tall premaxilla (frontmost upper jaw bone), which made the tip of the snout very blunt,
3920-473: Is parallel to the ground, the entire skull is nearly horizontal. This contrasts with many other theropods, where the head was more strongly downturned when in the alert position. The lateral canal is also significantly longer in Majungasaurus than in its more basal relative Ceratosaurus , indicating a greater sensitivity to side-to-side motions of the head. A 2007 report described pathologies in
4032-453: Is perhaps most distinctive for its skull ornamentation, including the swollen and fused nasals and the frontal horn. Other ceratosaurs, including Carnotaurus , Rajasaurus , and Ceratosaurus itself bore crests on the head. These structures are likely to have played a role in intraspecific competition , although their exact function within that context is unknown. The hollow cavity inside the frontal horn of Majungasaurus would have weakened
4144-759: 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
4256-607: The Northern Hemisphere , all of them being found in North America and Asia. They were all bipedal , herbivorous / omnivorous animals with thick skulls. Skulls can be domed, flat, or wedge-shaped depending on the species, and are all heavily ossified. The domes were often surrounded by nodes and/or spikes. Partial skeletons have been found of several pachycephalosaur species, but to date no complete skeletons have been discovered. Often isolated skull fragments are
4368-771: The State University of New York at Stony Brook and the University of Antananarivo began the Mahajanga Basin Project, a series of expeditions to examine the fossils and geology of the Late Cretaceous sediments near the village of Berivotra, in Mahajanga Province. Among these scientists was paleontologist David W. Krause of Stony Brook. The first expedition turned up hundreds of theropod teeth identical to those of Majungasaurus , some of which were attached to an isolated premaxilla that
4480-407: The center of gravity over the hips. Although the cervical ( neck ) vertebrae had numerous cavities and excavations ( pleurocoels ) to reduce their weight, they were robust, with exaggerated muscle attachment sites and ribs that interlocked for strength. Ossified tendons attached to the cervical ribs, giving them a forked appearance, as seen in Carnotaurus . All of these features resulted in
4592-400: The maxilla of the upper jaw and the dentary of the lower jaw, more than in any other abelisaurid except Rugops . The postcranial skeleton of Majungasaurus closely resembles those of Carnotaurus and Aucasaurus , the only other abelisaurid genera for which complete skeletal material is known. Majungasaurus was bipedal, with a long tail to balance out the head and torso, putting
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4704-622: The pelvis and the tip of the tail are still unknown. This fieldwork culminated in a 2007 monograph consisting of seven scientific papers on all aspects of the animal's biology, published in the Society of Vertebrate Paleontology Memoirs . The papers are in English, although each has an abstract written in Malagasy . In this volume, the dentary described by Lavocat was re-evaluated and determined to be diagnostic for this species. Therefore,
4816-501: 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)
4928-1031: 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"):
5040-495: 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
5152-556: 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, Majungatholus Majungasaurus ( / m ə ˌ dʒ ʌ ŋ ɡ ə ˈ s ɔː r ə s / ; lit. ' Mahajanga lizard ' )
5264-471: The analysis published by Woodruff and colleagues in 2023 is below. Wannanosaurus Hanssuesia Colepiocephale Stegoceras validum Stegoceras novomexicanum Goyocephale Homalocephale Tylocephale Foraminacephale Amtocephale Acrotholus Prenocephale Alaskacephale Stygimoloch Pachycephalosaurus S. goodwini S. lyonsi S. triregnum S. buchholtzae S. edmontonensis Below
5376-613: The animal's center of gravity. Thus Majungasaurus may have sacrificed speed for power. Majungasaurus tooth marks on Rapetosaurus bones confirm that it at least fed on these sauropods, whether or not it actually killed them. Although sauropods may have been the prey of choice for Majungasaurus , discoveries published in 2007 detail finds in Madagascar that indicate the presence of other Majungasaurus in their diet. Numerous bones of Majungasaurus have been discovered bearing tooth marks identical to those found on sauropod bones from
5488-415: The animals traditionally considered ' heterodontosaurids ' as a basal grouping within Pachycephalosauria, paraphyletic with respect to the traditional, dome-headed pachycephalosaurs. The same conclusion had previously been reached by George Olshevsky in 1991, who classified heterodontosaurids as basal pachycephalosaurs on the basis of perceived cranial kinesis , the presence of fanglike premaxillary teeth, and
5600-550: 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
5712-419: The augment when it was word-initial. In verbs with a preposition as a prefix, the augment is placed not at the start of the word, but between the preposition and the original verb. For example, προσ(-)βάλλω (I attack) goes to προσ έ βαλoν in the aorist. However compound verbs consisting of a prefix that is not a preposition retain the augment at the start of the word: αὐτο(-)μολῶ goes to ηὐ τομόλησα in
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#17327903659665824-475: The best-studied theropod dinosaurs from the Southern Hemisphere . It appears to be most closely related to abelisaurids from India rather than South America or continental Africa, a fact that has important biogeographical implications. Majungasaurus was the apex predator in its ecosystem , mainly preying on sauropods like Rapetosaurus , and is also one of the few dinosaurs for which there
5936-463: The body is never mixed with exhaled air laden with carbon dioxide . This method of respiration, while complicated, is highly efficient. The recognition of pneumatic foramina in Majungasaurus , besides providing an understanding of its respiratory biology, also has larger-scale implications for evolutionary biology. The split between the ceratosaur line, which led to Majungasaurus , and the tetanuran line, to which birds belong, occurred very early in
6048-450: The bones of Majungasaurus . Scientists examined the remains of at least 21 individuals and discovered four with noticeable pathologies. While pathology had been studied in large tetanuran theropods like allosaurids and tyrannosaurids , this was the first time an abelisauroid had been examined in this manner. No wounds were found on any skull elements, in contrast to tyrannosaurids where sometimes gruesome facial bites were common. One of
6160-479: The brain was very small relative to body size, but otherwise similar to many other non- coelurosaurian theropods, with a very conservative form closer to modern crocodilians than to birds. One difference between Majungasaurus and other theropods was its smaller flocculus , a region of the cerebellum that helps to coordinate movements of the eye with movements of the head. This suggests that Majungasaurus and other abelisaurids like Indosaurus , which also had
6272-438: 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
6384-413: The characteristic pits and grooves where claws and tendons would normally attach, and its finger bones were fused together, indicating that the hand was immobile. In 2012, a better specimen was described, showing that the lower arm was robust, though short, and that the hand contained four metatarsals and four, probably inflexible and very reduced, fingers, possibly with no claws. The minimum phalanx formula
6496-494: The contemporaneous marine environment. Besides Majungasaurus , fossil taxa recovered from the Maevarano include fish , frogs , lizards, snakes, seven distinct species of crocodylomorphs , five or six species of mammals , Vorona and several other birds, the possibly flighted dromaeosaurid Rahonavis , the noasaurid Masiakasaurus and two titanosaurian sauropods, including Rapetosaurus . Majungasaurus
6608-615: 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
6720-565: The distinctive dome. This can make taxonomic identification a difficult task, as the classification of genera and species within Pachycephalosauria relies almost entirely on cranial characteristics. Consequently, improper species have historically been appointed to the clade. For instance, Majungatholus , once thought to be a pachycephalosaur, is now recognized as a specimen of the abelisaurid theropod Majungasaurus , and Yaverlandia , another dinosaur initially described as
6832-430: The form of blunt horns and nodes. Many species are only known from skull fragments, and a complete pachycephalosaur skeleton is yet to be found. Members of Pachycephalosauria characteristically have an unusually domed head reminiscent of the earlier Protopyknosia in an example of convergent evolution . Most pachycephalosaurid remains are not complete, usually consisting of portions of the frontoparietal bone that forms
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#17327903659666944-673: 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
7056-463: The ground. Majungasaurus is classified as a member of the theropod clade Abelisauridae, which is considered a family in Linnaean taxonomy . Along with the family Noasauridae , abelisaurids are included in the superfamily Abelisauroidea , which is in turn a subdivision of the infraorder Ceratosauria . Abelisaurids are known for their tall skulls with blunt snouts, extensive sculpturing on
7168-566: The ground. Mallon et al. (2013) examined herbivore coexistence on the island continent of Laramidia , during the Late Cretaceous and concluded that pachycephalosaurids were generally restricted to feeding on vegetation at, or below, the height of 1 meter. They exhibit heterodonty , having different tooth morphology between the premaxillary teeth and maxillary teeth. Front teeth are small and peg-like with an ovular cross section and were most likely used for grabbing food. In some species,
7280-541: The half of the bone closest to the nostrils. A distinctive dome-like horn protruded from the fused frontal bones on top of the skull as well. In life, these structures would have been covered with some sort of integument , possibly made of keratin . Computed tomography (CT scanning) of the skull shows that both the nasal structure and the frontal horn contained hollow sinus cavities, perhaps to reduce weight. The teeth were typical of abelisaurids in having short crowns , although Majungasaurus bore seventeen teeth in both
7392-578: The head and neck for a bite-and-hold type of attack might have been very useful against sauropods, which would have been tremendously powerful animals. This hypothesis may also be supported by the hindlegs of Majungasaurus , which were short and stocky, as opposed to the longer and more slender legs of most other theropods. While Majungasaurus would not have moved as fast as other similar-sized theropods, it would have had no trouble keeping up with slow-moving sauropods. The robust hindlimb bones suggest very powerful legs, and their shorter length would have lowered
7504-524: The head, neck, and body be oriented in such a position. Instead, the cervical and anterior dorsal vertebrae of pachycephalosaurs show that the neck was carried in an "S"- or U-shaped curve. Also, the rounded shape of the skull would lessen the contacted surface area during head-butting, resulting in glancing blows. Other possibilities include flank-butting, defense against predators, or both. The relatively wide build of pachycephalosaurs (which would protect vital internal organs from harm during flank-butting) and
7616-540: The head-butting hypothesis, including Galton (1970) and Sues (1978) . In this hypothesis, pachycephalosaurs rammed each other head-on, as do modern-day bighorn sheep and musk oxen . Anatomical evidence for combative behavior includes vertebral articulations providing spinal rigidity, and the shape of the back indicating strong neck musculature. It has been suggested that a pachycephalosaur could make its head, neck, and body horizontally straight, in order to transmit stress during ramming. However, in no known dinosaur could
7728-561: 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
7840-476: 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
7952-430: The history of theropods. The avian respiratory system, present in both lines, must therefore have evolved before the split, and well before the evolution of birds themselves. This provides further evidence of the dinosaurian origin of birds . Computed tomography , also known as CT scanning, of a complete Majungasaurus skull (FMNH PR 2100) allowed a rough reconstruction of its brain and inner ear structure. Overall,
8064-402: The infection may have come first and led to the end of the tail becoming necrotic and falling off. This is the first example of tail truncation known in a non-avian theropod dinosaur. The small number of specimens preserved with pathologies in Majungasaurus suggest that the multitude of injuries that occurred were obtained over the course of the lives of the individuals studied. Furthermore,
8176-508: The last premaxillary tooth was enlarged and canine-like. Back teeth are small and triangular with denticles on the front and back of the crown, used for mouth processing. In species in which the dentary has been found, mandibular teeth are similar in size and shape to those in the upper jaw. Wear patterns on the teeth vary by species, indicating a range of food preferences which could include seeds, stems, leaves, fruits, and possibly insects. A very wide rib cage and large gut cavity extending all
8288-483: The likelihood of additional maladies and injuries due to functional impairment or compromised immune systems in individuals once an initial injury had occurred. Majungasaurus , being known from many well-preserved specimens of different ages, is well studied in regards to its growth and development. Throughout ontogeny, the skull of Majungasaurus (more specifically, the jugal , postorbital , and quadratojugal ) seems to have become taller and more robust; additionally,
8400-399: The name Majungatholus was replaced by the older name Majungasaurus . Although the monograph is comprehensive, the editors noted that it describes only material recovered from 1993 through 2001. A significant quantity of specimens, some very complete, were excavated in 2003 and 2005 and await preparation and description in future publications. The dentary was made the neotype specimen after
8512-451: The neck vertebrae and ribs are hollowed out by the cervical air sac, the upper back vertebrae by the lung , and the lower back and sacral (hip) vertebrae by the abdominal air sac. Similar features in Majungasaurus vertebrae imply the presence of these air sacs. These air sacs may have allowed for a basic form of avian-style 'flow-through ventilation,' where air flow through the lungs is one-way, so that oxygen -rich air inhaled from outside
8624-475: The new genus Majungasaurus , using an older spelling of Mahajanga as well as the Greek word σαυρος sauros (meaning "lizard"). Hans-Dieter Sues and Philippe Taquet described a dome-shaped skull fragment (MNHN.MAJ 4) as a new genus of pachycephalosaur ( Majungatholus atopus ) in 1979. This was the first report of a pachycephalosaur in the Southern Hemisphere. In 1993, scientists from
8736-648: The next 100 years, many of which were deposited in the Muséum National d'Histoire Naturelle in Paris . In 1955, René Lavocat described a theropod dentary ( MNHN .MAJ 1) with teeth from the Maevarano Formation in the same region where the original material was found. The teeth matched those first described by Depéret, but the strongly curved jaw bone was very different from both Megalosaurus and Dryptosaurus . Based on this dentary, Lavocat created
8848-617: The occurrence of periodic debris flows through these channels at the beginning of the wet season, burying the carcasses of organisms killed during the preceding dry season and providing for their exceptional preservation as fossils. Sea levels in the area were rising throughout the Maastrichtian, and would continue to do so into the Paleocene Epoch , so Majungasaurus may have roamed coastal environments like tidal flats as well. The neighboring Berivotra Formation represents
8960-508: 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
9072-493: The only bones that are found. Candidates for the earliest-known pachycephalosaur include Ferganocephale adenticulatum from Middle Jurassic Period strata of Kyrgyzstan and Stenopelix valdensis from Early Cretaceous strata of Germany , although R.M. Sullivan has doubted that either of these species are pachycephalosaurs. Albalophosaurus from the Early Cretaceous strata of Japan might also represent
9184-571: The original diagnosis of Pachycephalosauridae centered around "flat to dome-like" skulls, so the flat-headed forms should be included in the family. In a paper published in 2003, Thomas E. Williamson and Thomas D. Carr discovered a clade of the Pachycephalosauridae that was a sister taxa to the genus Stegoceras , made up of "all other dome-headed pachycephalosaurians; this was referred to as Pachycephalosaurinae Phylogenetic analyses by many authors have found Pachycephalosauria to be
9296-470: The ossification of a ligament running either between the neural spines ( interspinal ligament ) or along their tops ( supraspinal ligament ). The most serious pathology discovered was in a series of five large tail vertebrae. The first two vertebrae showed only minor abnormalities with the exception of a large groove that extended along the left side of both bones. However, the next three vertebrae were completely fused together at many different points, forming
9408-488: The ossification was not determined. Hypervitaminosis A and bone spurs were ruled out, and an osteoma (benign bone tumor) was deemed unlikely. Another specimen, a small caudal (tail) vertebra, was also found to have an abnormal growth, this time on the top of its neural spine , which projects upwards from the vertebrae, allowing muscle attachment. Similar growths from the neural spine have been found in specimens of Allosaurus and Masiakasaurus , probably resulting from
9520-411: The outer surfaces of the facial bones ( convergent with carcharodontosaurids ), very reduced ( atrophied ) forelimbs (convergent with tyrannosaurids ), and stocky hindlimb proportions, among other features. As with many dinosaur families, the systematics ( evolutionary relationships) within the family Abelisauridae are confused. Several cladistic studies have indicated that Majungasaurus shares
9632-487: 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
9744-443: The possibility of separate clades of abelisaurids in western and eastern Gondwana . A cladogram by Tortosa et al. 2013 places Majungasaurus in a new subfamily, Majungasaurinae . A simplified version showing the taxa within the group is shown below. Pourcieux abelisaurid Arcovenator [REDACTED] Majungasaurus [REDACTED] Indosaurus Rahiolisaurus Rajasaurus [REDACTED] Majungasaurus
9856-414: The prey is subdued. Majungasaurus had an even broader snout than other abelisaurids, and other aspects of its anatomy may also support the bite-and-hold hypothesis. The neck was strengthened, with robust vertebrae, interlocking ribs and ossified tendons, as well as reinforced muscle attachment sites on the vertebrae and the back of the skull. These muscles would have been able to hold the head steady despite
9968-429: The prominent diastema present in many genera. The Pachycephalosauria was first named as a suborder of the order Ornithischia by Maryańska & Osmólska (1974) . They included within it only one family , the Pachycephalosauridae. Later researchers, such as Michael Benton, have ranked it as an infraorder of a suborder Cerapoda , which unites the ceratopsians and ornithopods . In 2006, Robert Sullivan published
10080-463: The remains of their rivals, which may suggest similar behavior in Majungasaurus and other theropods. Scientists have reconstructed the respiratory system of Majungasaurus based on a superbly preserved series of vertebrae ( UA 8678) recovered from the Maevarano Formation. Most of these vertebrae and some of the ribs contained cavities (pneumatic foramina ) that may have resulted from the infiltration of avian -style lungs and air sacs . In birds,
10192-480: The same localities. These marks have the same spacing as teeth in Majungasaurus jaws, are of the same size as Majungasaurus teeth, and contain smaller notches consistent with the serrations on those teeth. As Majungasaurus is the only large theropod known from the area, the simplest explanation is that it was feeding on other members of its own species. Suggestions that the Triassic Coelophysis
10304-417: The skull bones became more fused and the eye sockets became proportionally smaller. This indicates a shift in dietary preferences between juveniles and adults. Research by Michael D'Emic et al indicates that it was among the slowest-growing theropods. Based on studies of the lines of arrested growth in several bones, it was found that Majungasaurus took twenty years to reach maturity, which may have been
10416-403: The skull. Further fieldwork over the next decade turned up a series of less complete skulls, as well as dozens of partial skeletons of individuals ranging from juveniles to adults. Project members also collected hundreds of isolated bones and thousands of shed Majungasaurus teeth. Taken together, these remains represent nearly all the bones of the skeleton, although most of the forelimbs, most of
10528-410: The small number of injured Majungasaurus specimens observed amongst those recovered indicates most well preserved individuals generally lack observable pathologies, while a few select individuals were shown to have possessed multiple pathologies, a general pattern also noted in other large, nonavian theropods. Such patterns may be the result of a snowball effect where one injury or an infection increased
10640-405: The specimens was a phalanx (toe bone) of the foot, which had apparently been broken and subsequently healed. Most of the pathologies occurred on the vertebrae. For example, a dorsal (back) vertebra from a juvenile animal showed an exostosis (bony growth) on its underside. The growth probably resulted from the conversion of cartilage or a ligament to bone during development, but the cause of
10752-475: The squamosal horns of the Stygimoloch (which would have been used to great effect during flank-butting) add credence to the flank-butting hypothesis. A histological study conducted by Goodwin & Horner (2004) argued against the battering ram hypothesis. They argued that the dome was "an ephemeral ontogenetic stage", the spongy bone structure could not sustain the blows of combat, and the radial pattern
10864-636: The structure and probably precluded its use in direct physical combat, although the horn may have served a display purpose. While there is variation in the ornamentation of Majungasaurus individuals, there is no evidence for sexual dimorphism . Scientists have suggested that the unique skull shape of Majungasaurus and other abelisaurids indicate different predatory habits than other theropods. Whereas most theropods were characterized by long, low skulls of narrow width, abelisaurid skulls were taller and wider, and often shorter in length as well. The narrow skulls of other theropods were well equipped to withstand
10976-486: The struggles of its prey. Abelisaurid skulls were also strengthened in many areas by bone mineralized out of the skin , creating the characteristic rough texture of the bones. This is particularly true of Majungasaurus , where the nasal bones were fused and thickened for strength. On the other hand, the lower jaw of Majungasaurus sported a large fenestra (opening) on each side, as seen in other ceratosaurs, as well as synovial joints between certain bones that allowed
11088-517: 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
11200-517: The theropod replaced its teeth anywhere from 2 to 13 times faster than other theropods, replacing the entire set within a span of two months. Gnawing on bone may have been a significant reason for such rapid tooth replacement. Majungasaurus was the largest predator in its environment, while the only known large herbivores at the time were sauropods like Rapetosaurus . Scientists have suggested that Majungasaurus , and perhaps other abelisaurids, specialized on hunting sauropods. Adaptations to strengthen
11312-666: The time was a wastebasket taxon containing any number of unrelated large theropods, as the new species M. crenatissimus . This name is derived from the Latin word crenatus ("notched") and the suffix -issimus ("most"), in reference to the numerous serrations on both front and rear edges of the teeth. Depéret later reassigned the species to the North American genus Dryptosaurus , another poorly known taxon. Numerous fragmentary remains from Mahajanga Province in northwestern Madagascar were recovered by French collectors over
11424-510: The underlying Masorobe Member and the overlying Miadana Member. While these sediments have not been dated radiometrically , evidence from biostratigraphy and paleomagnetism suggest that they were deposited during the Maastrichtian stage , which lasted from 70 to 66 Ma (million years ago). Majungasaurus teeth are found up until the very end of the Maastrichtian, when all non-avian dinosaurs became extinct . Then as now, Madagascar
11536-500: The vertical stress of a powerful bite, but not as good at withstanding torsion (twisting). In comparison to modern mammalian predators , most theropods may have used a strategy similar in some ways to that of long- and narrow-snouted canids , with the delivery of many bites weakening the prey animal. Abelisaurids, especially Majungasaurus , may instead have been adapted for a feeding strategy more similar to modern felids , with short and broad snouts, that bite once and hold on until
11648-556: The very rough texture and thickened bone on the top of its snout, and the single rounded horn on the roof of its skull, which was originally mistaken for the dome of a pachycephalosaur . It also had more teeth in both upper and lower jaws than most abelisaurids. The genus is one of the first abelisaurs to be discovered, first being found in 1896 (although it was thought to be a species of Megalosaurus ) and being named in 1955. Known from several well-preserved skulls and abundant skeletal material, Majungasaurus has recently become one of
11760-482: The way to the base of the tail suggests the use of fermentation to digest food. The adaptive significance of the skull dome has been heavily debated. The popular hypothesis among the general public that the skull was used in head-butting, as sort of a dinosaurian battering ram, was first proposed by Colbert (1955) , p. . This view was popularized in the 1956 science fiction story " A Gun for Dinosaur " by L. Sprague de Camp . Many paleontologists have since argued for
11872-403: Was 1-2-1-1. Like other abelisaurids, the hindlimbs were stocky and short compared to body length. The tibia (lower leg bone) of Majungasaurus was even stockier than that of its relative Carnotaurus , with a prominent crest on the knee. The astragalus and calcaneum (ankle bones) were fused together, and the feet bore three functional digits, with a smaller first digit that did not contact
11984-480: 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 ,
12096-519: Was a cannibal have been recently disproven, leaving Majungasaurus as the only non-avian theropod with confirmed cannibalistic tendencies, although there is some evidence that cannibalism may have occurred in other species as well. It is unknown if Majungasaurus actively hunted their own kind or only scavenged their carcasses. However, some researchers have noted that modern Komodo monitors sometimes kill each other when competing for access to carcasses. The lizards will then proceed to cannibalize
12208-402: Was also typical of the family. However, the skull of Majungasaurus was markedly wider than in other abelisaurids. All abelisaurids had a rough, sculptured texture on the outside faces of the skull bones, and Majungasaurus was no exception. This was carried to an extreme on the nasal bones of Majungasaurus , which were extremely thick and fused together, with a low central ridge running along
12320-491: Was an island, having separated from the Indian subcontinent less than 20 million years earlier. It was drifting northwards but still 10 to 15 degrees more southerly in latitude than it is today. The prevailing climate of the time was semi-arid, with pronounced seasonality in temperature and rainfall. Majungasaurus inhabited a coastal flood plain cut by many sandy river channels . Strong geological evidence suggests
12432-456: Was described in 1996. The following seven expeditions would turn up tens of thousands of fossils, many of which belonged to species new to science. The Mahajanga Basin Project claims credit for quintupling the known diversity of fossil taxa in the region. Fieldwork in 1996 turned up a spectacularly complete theropod skull preserved in exquisite detail ( FMNH PR 2100). On top of this skull was
12544-430: Was simply an effect of rapid growth. Later biomechanical analyses by Snively & Cox (2008) and Snively & Theodor (2011) concluded, however, that the domes could withstand combat stresses. Lehman (2010) argued that the growth patterns discussed by Goodwin and Horner are not inconsistent with head-butting behavior. Goodwin & Horner (2004) instead argued that the dome functioned for species recognition. There
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