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Compsognathus

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60-465: Compsognathus ( / k ɒ m p ˈ s ɒ ɡ n ə θ ə s / ; Greek kompsos /κομψός; "elegant", "refined" or "dainty", and gnathos /γνάθος; "jaw") is a genus of small, bipedal , carnivorous theropod dinosaur . Members of its single species Compsognathus longipes could grow to around the size of a chicken . They lived about 150  million years ago , during the Tithonian age of

120-543: 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

180-417: A century, Compsognathus longipes was the only well-known small theropod species. This led to comparisons with Archaeopteryx and to suggestions of an especially close relationship with birds. In fact, Compsognathus , rather than Archaeopteryx , piqued Huxley's interest in the origin of birds. The two animals share similarities in shape and proportions, so many in fact that two specimens of Archaeopteryx ,

240-477: 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

300-419: 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

360-631: A new clade of dinosaurs, the Symphypoda, which also contained Ornithotarsus (today classified as Hadrosaurus ). Later, both genera were found to belong to other groups of Cope's classification of dinosaurs: Compsognathus to the Gonipoda (equivalent to Theropoda , in which it is now classified), and Ornithotarsus to the Orthopoda (equivalent to Ornithischia ). Huxley, in 1870, rejected Cope's dinosaur classification scheme, and instead proposed

420-562: A patch of fossilized skin from the tail and hindlimb of the possible relative Juravenator starki shows mainly scales, though there is some indication that simple feathers were also present in the preserved areas. This may mean that a feather covering was not ubiquitous in this group of dinosaurs, or maybe that some species had fewer feathers than others. Originally classified as a lizard, the dinosaurian affinities of Compsognathus were first noted by Gegenbaur, Cope, and Huxley between 1863 and 1868. Cope, in 1870, classified Compsognathus within

480-550: 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

540-608: 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

600-630: 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

660-510: 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

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720-570: 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

780-418: 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

840-705: Is an alternative phylogeny, published by Cau in 2024, with both specimens in bold. Siamraptor Siamotyrannus Streptospondylus Xuanhanosaurus Poekilopleuron Piveteausaurus Piatnitzkysaurus Marshosaurus Leshansaurus Eustreptospondylus Condorraptor Asfaltovenator Sciurumimus Nedcolbertia Magnosaurus Duriavenator Afrovenator Compsognathus longipes Compsognathus corallestris Torvosaurus tanneri Torvosaurus gurneyi Megalosaurus Scipionyx Wiehenvenator Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes

900-666: 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

960-448: 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

1020-464: Is the first theropod dinosaur known from a reasonably complete fossil skeleton. Until the 1990s, it was the smallest-known non- avialan dinosaur, with the preceding centuries incorrectly labelling them as the closest relative of Archaeopteryx . Compsognathus is known from two almost complete skeletons. The German specimen (specimen number BSP AS I 563) stems from limestone deposits in Bavaria and

1080-587: The Compsognathus specimen, was acquired by the paleontological state collection in Munich . Both the year of discovery and the exact locality of the German specimen are unknown, possibly because Oberndorfer did not reveal details of the discovery to prevent other collectors from exploiting the locality; later authors have suggested that the German specimen was probably discovered during the 1850s. Weathering of

1140-759: 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

1200-501: 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)

1260-514: The "Eichstätt" and the "Solnhofen", were for a time misidentified as those of Compsognathus . Many other types of theropod dinosaurs, such as maniraptorans , are now known to have been more closely related to birds. Below is a simplified cladogram placing Compsognathus in Compsognathidae by Senter et al. in 2012. Sinocalliopteryx Huaxiagnathus Sinosauropteryx Compsognathus Juravenator Scipionyx Here

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1320-1031: 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"):

1380-495: 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

1440-550: 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

1500-419: 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

1560-438: 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

1620-466: The compsognathids to be an example of convergent evolution . The position of Compsognathus and its relatives within the coelurosaur group is uncertain. Some, such as theropod expert Thomas Holtz Jr. and co-authors Ralph Molnar and Phil Currie in the landmark 2004 text Dinosauria , hold the family as the most basal of the coelurosaurs, while others as part of the Maniraptora . For almost

1680-615: 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

1740-566: The first two digits, leading to the suggestion that Compsognathus bore only two functional digits, with the third metacarpal being extremely slender and reduced. Study of the French specimen indicated that the third digit bore at least one or two small phalanges. However, there remains no evidence for an ungual phalanx on the third digit, so the digit may have been reduced and non-functional. Some relatives of Compsognathus , namely Sinosauropteryx and Sinocalliopteryx , have been preserved with

1800-471: The foot did probably not belong to Compsognathus itself but to a closely related genus. Ostrom, in his 1978 monography, questioned the attribution of this fossil to Compsognathus once more. Jens Zinke, in 1998, assigned forty-nine isolated teeth from the Guimarota coal mine of Portugal to the genus. Zinke found that these teeth are not identical to those of Compsognathus longipes , having serrations on

1860-673: 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

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1920-410: The front edge, and thus labeled the teeth as Compsognathus sp. (of unknown species). For decades, Compsognathus was known as the smallest known non-avian dinosaur , although some dinosaurs discovered later, such as Mahakala and Microraptor , were even smaller. The German specimen was estimated to be 70–75 cm (28–30 in) and 89 cm (35 in) in length by separate authors, while

1980-401: The frontmost teeth of the upper and lower jaws were unserrated , while those further back had fine serrations on their rear edges. In the German specimen, the crowns were around two times higher than wide in the front of the jaws but diminished in height further back, with the last tooth about as high as wide. The German specimen also shows a diastema (tooth gap) behind the first three teeth of

2040-561: 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

2100-476: 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

2160-499: The larger French specimen was estimated at 1.25 m (4 ft 1 in) and 1.4 m (4 ft 7 in) in length. The height at the hip has been estimated at 20 cm (7.9 in) for the German specimen and at 29 cm (11 in) for the French specimen. The German specimen was estimated to have weighed 0.32 kg (0.71 lb) and 0.58 kg (1.3 lb), and the French specimen 2.5 kg (5.5 lb) and 3.5 kg (7.7 lb). Compared to other compsognathids,

2220-555: The larger French specimen would have been similar in size to larger Sinosauropteryx specimens, but smaller than Huaxiagnathus and Mirischia . Compsognathus were small, bipedal animals with long hind legs and longer tails, which they used for balance during locomotion. The forelimbs were smaller than the hindlimbs. The hand bore two large, clawed digits and a third, smaller digit that may have been non-functional. Their delicate skulls were narrow and long, with tapered snouts. The skull had five pairs of fenestrae (skull openings),

2280-449: The largest of which was for the orbit (eye socket), with the eyes being larger in proportion to the rest of the skull. The lower jaw was slender and had no mandibular fenestra, a hole in the side of the lower jawbone commonly seen in archosaurs . The teeth were small and pointed, suited for its diet of small vertebrates and possibly other small animals, such as insects. The German specimen had three teeth in each premaxilla (front bone of

2340-640: The late Jurassic period , in what is now Europe . Paleontologists have found two well-preserved fossils , one in Germany in the 1850s and the second in France more than a century later. Today, C. longipes is the only recognized species, although the larger specimen discovered in France in the 1970s was once thought to belong to a separate species and named C. corallestris . Many presentations still describe Compsognathus as "chicken-sized" dinosaurs because of

2400-427: The lower jaw), 15 or 16 teeth in each maxilla, and 18 teeth in the lower jaw. The French specimen had more teeth, including four in each premaxilla, 17 or 18 in the maxilla, and at least 21 teeth in the dentary. Compsognathids were unique among theropods in having tooth crowns that curved backwards at two thirds of their height, while their mid-parts were straight; also, the crowns had expanded bases. In Compsognathus ,

2460-601: The new clade Ornithoscelida , in which he included the Dinosauria (comprising several forms now considered as ornithischians) and another new clade, the Compsognatha, which contained Compsognathus as the only member. Later, these groups fell into disuse, although a resurrection of the Ornithoscelida was proposed in 2017. The group Compsognatha was used for the last time by Marsh in a 1896 publication, where it

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2520-418: The new find under a separate species, Compsognathus corallestris . A more comprehensive description followed in the same year. According to these authors, the new species differed from the German species in its larger size and modified, flipper-like hand. Ostrom, Jean-Guy Michard and others have since relabeled it as another example of Compsognathus longipes . In 1984, George Callison and Helen Quimby identified

2580-505: The newly discovered French specimen in 1978, making Compsognathus one of the best-known small theropods at that time. He also concluded that the German specimen likely belongs to an immature individual. The larger French specimen (Y85R M4M) was discovered in around 1971 in the Portlandian lithographic limestone of Canjuers near Nice . It dates to the lower Tithonian, as indicated by ammonite index fossils . As Solnhofen, Canjures

2640-508: 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

2700-487: 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

2760-417: The premaxilla. As such a gap was not present in the French specimen, Peyer suggested that additional teeth were possibly present in this region the German specimen. The number of digits on the hand of Compsognathus has been a source of debate. For much of its history, Compsognathus was typically depicted with three digits, as is typical for theropods. However, the type specimen only preserved phalanges from

2820-562: The region was part of the Solnhofen archipelago. The limestone of the area, the Solnhofen limestone , had been quarried for centuries, and yielded such well-preserved fossils as Archaeopteryx with feather impressions and pterosaurs with imprints of their wing membranes. In two publications in 1868 and 1870, Thomas Huxley , a major proponent of Charles Darwin 's theory of evolution, compared Compsognathus with Archaeopteryx , which

2880-406: The remains of simple feathers covering the body like fur, prompting some scientists to suggest that Compsognathus might have been feathered in a similar way. Consequently, many depictions of Compsognathus show them with coverings of downy proto-feathers. However, no feathers or feather-like covering have been preserved with Compsognathus fossils, in contrast to Archaeopteryx , which are found in

2940-402: The right ankle from the slab for illustration and study; this removed part got lost since. Although Baur published a detailed study of the ankle in 1882, which is now the only available source of information of this part of the skeleton, his reconstruction was later found to be inconsistent with corresponding impressions on the slab. John Ostrom thoroughly described the German specimen as well as

3000-685: The same sediments. Karin Peyer, in 2006, reported skin impressions preserved on the side of the tail starting at the 13th tail vertebra. The impressions showed small bumpy tubercles, similar to the scales found on the tail and hind legs of Juravenator . Additional scales had in 1901 been reported by Von Huene, in the abdominal region of the German Compsognathus , but Ostrom subsequently disproved this interpretation; in 2012 they were by Achim Reisdorf seen as plaques of adipocere , corpse wax. Like Compsognathus , and unlike Sinosauropteryx ,

3060-460: The size of the German specimen, which is now believed to be a juvenile. Compsognathus longipes is one of the few dinosaur species whose diet is known with certainty: the remains of small, agile lizards are preserved in the bellies of both specimens. Teeth discovered in Portugal may be further fossil remains of the genus. Although not recognized as such at the time of its discovery, Compsognathus

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3120-527: The slab on which the fossil is preserved indicates that it was collected from a pile of waste rock left behind by quarrying. The specimen either stems from Jachenhausen or the region Riedenburg–Kehlheim. All possible localities are part of lagoonal deposits of the Painten Formation , and date to the latest part of the late Kimmeridgian or the earlier part of the early Tithonian . In the Jurassic,

3180-412: The smaller German specimen as a juvenile of the same species. Collector Heinrich Fischer had originally labeled a partial foot consisting of three metatarsals and a phalanx, from the Solnhofen area, as belonging to Compsognathus longipes . This identification was rejected by Wilhelm Dames, when he described the specimen for the first time in 1884. Friedrich von Huene , in 1925 and 1932, also found that

3240-517: 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

3300-480: 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 ,

3360-607: Was considered the earliest known bird. Following earlier suggestions by Carl Gegenbaur and Edward Drinker Cope , Huxley found that Archaeopteryx was closely similar to Compsognathus , and referred to the latter as a "bird-like reptile". He concluded that birds must have evolved from dinosaurs, an assessment that established Compsognathus as one of the most widely known dinosaurs. The specimen has since been studied by many prominent paleontologists, including Othniel Charles Marsh , who visited Munich in 1881. The German paleontologist J.G. Baur, who worked as an assistant of Marsh, removed

3420-649: Was famous for its limestone plates, which were quarried and sold under the name "dalles de Provence". The specimen was originally part of a large private fossil collection of Louis Ghirardi, the owner of the Canjures quarries. The collection, including the Compsognathus specimen, was sold to the National Museum of Natural History in Paris in 1983. Alain Bidar and Gérard Thomel, in a brief 1972 description, announced

3480-477: Was part of the collection of the physician and fossil collector Joseph Oberndorfer. Oberndorfer lent the specimen to paleontologist Johann A. Wagner , who published a brief discussion in 1859, where he coined the name Compsognathus longipes . Wagner did not recognise Compsognathus as a dinosaur, but instead described it as one of the "most curious forms among the lizards". He published a more detailed description in 1861. In 1866, Oberndorfer's collection, including

3540-430: Was the only member known, but in recent decades paleontologists have discovered several related genera. The clade includes Aristosuchus , Huaxiagnathus , Mirischia , Sinosauropteryx , and perhaps Juravenator and Scipionyx . At one time, Mononykus was proposed as a member of the family, but this was rejected by Chen and coauthors in a 1998 paper; they considered the similarities between Mononykus and

3600-490: Was treated as a suborder of Theropoda. In the same publication, Marsh erected the new family Compsognathidae . Friedrich von Huene, in 1914, erected the new infraorder Coelurosauria , which includes the Compsognathidae amongst other families of small theropods; this classification remained in use since. The Compsognathidae are a group of mostly small dinosaurs from the late Jurassic and early Cretaceous periods of China, Europe and South America. For many years, Compsognathus

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