191-517: Tanystropheus ( ‹See Tfd› Greek : τανυ ~ 'long' + στροφευς 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs . Tanystropheus
382-604: A nomen dubium possibly synonymous with T. hydroides . Over 500 " Tanystropheus conspicuus " specimens have been recovered from a Lower Keuper bonebed near the Silesian village of Miedary . This is the largest known concentration of Tanystropheus fossils, more than double the number collected from Monte San Giorgio. Though the Miedary specimens are individually limited to isolated postcranial bones, they are preserved in three dimensions and show great potential for elucidating
573-436: A Tanystropheus individual of that length, the weight estimate varied between 32.9 kg (72.5 lbs) and 74.8 kg (164.9 lbs), depending on the volume estimation method. This was significantly lighter than crocodiles of the same length, and more similar to large lizards. The skull of Tanystropheus longobardicus is roughly triangular when seen from the side and top, narrowing towards the snout. Each premaxilla (the toothed bone at
764-566: A junior synonym of T. longobardicus . A 2019 revision of Tanystropheus found that T. longobardicus and T. antiquus were the only valid species in the genus . Tanystropheus specimens from Monte San Giorgio have long been segregated into two morphotypes based on their tooth structure. Smaller specimens bear tricuspid teeth at the back of the jaw while larger specimens have a set of single-pointed fangs. The two morphotypes were originally considered to represent juvenile and adult specimens of T. longobardicus , though many studies have supported
955-692: A nomen dubium . Several more von Huene species, including " Procerosaurus cruralis ", " Thecodontosaurus latespinatus ", and " Thecodontosaurus primus ", have been reconsidered as indeterminate material of Tanystropheus or other archosauromorphs . One of Von Huene's species appears to be valid: T. antiquus , from the Gogolin Formation of Poland, was based on cervical vertebrae which were proportionally shorter than those of other Tanystropheus species. Long considered destroyed in World War II, several T. antiquus fossils were rediscovered in
1146-473: A European species from the latest part of the Early Triassic (late Olenekian stage). T. antiquus had a proportionally shorter neck than other Tanystropheus species, so some paleontologists consider that T. antiquus deserves a separate genus, Protanystropheus . The lifestyle of Tanystropheus has been the subject of much debate. Tanystropheus is unknown from drier environments and its neck
1337-577: A basis for coinages: anthropology , photography , telephony , isomer , biomechanics , cinematography , etc. Together with Latin words , they form the foundation of international scientific and technical vocabulary ; for example, all words ending in -logy ('discourse'). There are many English words of Greek origin . Greek is an independent branch of the Indo-European language family. The ancient language most closely related to it may be ancient Macedonian , which, by most accounts,
1528-421: A broad flat body and a short tail. Their limbs had evolved into four long flippers, which were powered by strong muscles attached to wide bony plates formed by the shoulder girdle and the pelvis. The flippers made a flying movement through the water. Plesiosaurs breathed air, and bore live young; there are indications that they were warm-blooded. Plesiosaurs showed two main morphological types. Some species, with
1719-424: A broad furrow on the underside. The parietals are strongly modified in T. hydroides . They are fused into a single X-shaped bone, somewhat resembling the parietals of erythrosuchids . This shape may have resulted from fusion between the parietals' anterolateral processes (front branches) and the postfrontals, which are separate bones in T. longobardicus but not apparent in T. hydroides . A prominent pineal foramen
1910-511: A cervical count of 13, Tanystropheus acquired four additional elongated cervicals in the front half of the neck, in addition to a stout vertebra which shifted from the dorsal series into the base of the neck, transforming into the 13th cervical. Tanystropheids are unusual among reptiles in that they acquire their long necks without prolonged somitogenesis (an increase in the overall number of presacral vertebrae during early development). Instead, their overall number of presacral vertebrae remains at
2101-426: A constant count of 25, the same as their shorter-necked ancestors. This would require a shift in regionalization, encouraging the development of new cervical vertebrae rather than dorsals. There are 12 dorsal (torso) vertebrae . This count is very low among early archosauromorphs: Protorosaurus has up to 19, Prolacerta has 18, and Macrocnemus has 17. Tanystropheus' s dorsals are smaller and less specialized than
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#17327802173782292-519: A crocodile or dolphin. The specimen is today on display at the Natural History Museum , and its inventory number is NHMUK PV R.1330 (formerly BMNH R.1330). It is the earliest discovered more or less complete fossil reptile skeleton in a museum collection. It can perhaps be referred to Plesiosaurus dolichodeirus . During the eighteenth century, the number of English plesiosaur discoveries rapidly increased, although these were all of
2483-558: A fairly stable set of consonantal contrasts . The main phonological changes occurred during the Hellenistic and Roman period (see Koine Greek phonology for details): In all its stages, the morphology of Greek shows an extensive set of productive derivational affixes , a limited but productive system of compounding and a rich inflectional system. Although its morphological categories have been fairly stable over time, morphological changes are present throughout, particularly in
2674-560: A faster, more convenient cursive writing style with the use of ink and quill . The Greek alphabet consists of 24 letters, each with an uppercase ( majuscule ) and lowercase ( minuscule ) form. The letter sigma has an additional lowercase form (ς) used in the final position of a word: In addition to the letters, the Greek alphabet features a number of diacritical signs : three different accent marks ( acute , grave , and circumflex ), originally denoting different shapes of pitch accent on
2865-465: A fatal hindrance to terrestrial locomotion. The hindlimbs and the base of the tail were large and muscular, capable of short bursts of active swimming in shallow water. Tanystropheus was most likely a piscivorous ambush predator : the narrow subtriangular skull of T. longobardicus is supplied with three-cusped teeth suited for holding onto slippery prey, while the broader skull of T. hydroides bears an interlocking set of large curved fangs similar to
3056-537: A few (very) basal forms have been discovered, the most derived Rhaeticosaurus . The subsequent evolution of the plesiosaurs is very contentious. The various cladistic analyses have not resulted in a consensus about the relationships between the main plesiosaurian subgroups. Traditionally, plesiosaurs have been divided into the long-necked Plesiosauroidea and the short-necked Pliosauroidea . However, modern research suggests that some generally long-necked groups might have had short-necked members. To avoid confusion between
3247-438: A few modern lizard species. Some individuals of T. longobardicus have tricuspid teeth along their entire maxilla, while in others up to seven maxillary teeth are single-cusped fangs similar to the premaxillary teeth. The front edge of each orbit (eye socket) is marked by two bones: the prefrontal and lacrimal . The prefrontal is tall and projects a low vertical ridge in front of the orbit. The small, sliver-shaped lacrimal
3438-540: A foreign language. It is also often stated that the historical changes have been relatively slight compared with some other languages. According to one estimation, " Homeric Greek is probably closer to Demotic than 12-century Middle English is to modern spoken English ". Greek is spoken today by at least 13 million people, principally in Greece and Cyprus along with a sizable Greek-speaking minority in Albania near
3629-540: A large skeleton of a plesiosaur was found in the continent of Antarctica, the oldest creature on the continent, and the first of its species in Antarctica. Not only has the number of field discoveries increased, but also, since the 1950s, plesiosaurs have been the subject of more extensive theoretical work. The methodology of cladistics has, for the first time, allowed the exact calculation of their evolutionary relationships. Several hypotheses have been published about
3820-448: A larger portion of the internal dentary than in T. longobardicus . In addition, the rear of the dentary overlaps a large portion of the surangular, rather than the surangular acting as the overlapping bone where they meet. The surangular internally bears a large fossa for the jaw's adductor (vertical biting) muscles, and a prominent surangular foramen is positioned in front of the jaw joint. The most recognisable feature of Tanystropheus
4011-537: A long and slender posterior process (rear branch) that projects under the rounded orbit. There are 15 teeth in the maxilla, increasing in size up to the eighth tooth, which is about as large as the premaxillary fangs. T.hydroides is not known to possess a septomaxilla , a neomorphic bone at the rear tip of the naris in some reptiles. The nasals are broad and plate-like, with a depressed central portion. The lacrimal and prefrontal, though incompletely known, were likely similar to those of T. longobardicus . T. hydroides has
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#17327802173784202-475: A loose, strongly overlapping connection to the ectopterygoids (linking bones between the pterygoid and maxilla). The epipterygoids (vertical bones in front of the braincase) are tall and flattened from the side. T. hydroides is a rare example of an early archosauromorph with a three-dimensionally preserved braincase. The basioccipital (rear lower component of the braincase) was small, with inset basitubera (vertical plates connecting to neck muscles) linked by
4393-417: A more basal type of reptile unrelated to Archosauromorpha. The following cladogram is from Dilkes (1998), a study with a small sample of "prolacertiforms" but closer resemblance to most analyses of the 2000s and 2010s: Greek language Greek ( Modern Greek : Ελληνικά , romanized : Elliniká , [eliniˈka] ; Ancient Greek : Ἑλληνική , romanized : Hellēnikḗ )
4584-435: A more comprehensive view of the species' anatomy. A small but well-preserved skull and neck, specimen PIMUZ T 3901, was found in the slightly younger Meride Limestone at Monte San Giorgio. Wild (1980) gave it a new species, T. meridensis , based on a set of skull and vertebral traits proposed to differ from T. longobardicus . Later reinvestigations failed to confirm the validity of these differences, rendering T. meridensis
4775-542: A more limited meaning. The term "plesiosaur" is properly used to refer to the Plesiosauria as a whole, but informally it is sometimes meant to indicate only the long-necked forms, the old Plesiosauroidea. Like other ancient marine reptiles, such as those in the clades Ichthyosauria and Mosasauria , the genera in Plesiosauria are not part of the clade Dinosauria . Skeletal elements of plesiosaurs are among
4966-518: A more or less fragmentary nature. Important collectors were the reverends William Mounsey and Baptist Noel Turner , active in the Vale of Belvoir , whose collections were in 1795 described by John Nicholls in the first part of his The History and Antiquities of the County of Leicestershire . One of Turner's partial plesiosaur skeletons is still preserved as specimen NHMUK PV R.45 (formerly BMNH R.45) in
5157-447: A new clade Neoplesiosauria, a node-based taxon that was defined by as " Plesiosaurus dolichodeirus , Pliosaurus brachydeirus , their most recent common ancestor and all of its descendants". The clade Neoplesiosauria very likely is materially identical to Plesiosauria sensu Druckenmiller & Russell, thus would designate exactly the same species, and the term was meant to be a replacement of this concept. Benson et al. (2012) found
5348-447: A pair of atlantal neural arches (prong-like upper components). There does not appear to be a proatlas , which slots between the atlas and skull in some other reptiles. The intercentrum and pleurocentrum are not fused to each other, unlike the single-part atlas of allokotosaurs . The tiny crescent-shaped intercentrum is overlain by a semicircular pleurocentrum, which acts as a base to the backswept neural arches. The axis (second cervical)
5539-483: A pair of segmented rods which intermingle at the midline. The two sacral (hip) vertebrae are low but robust, bridging over to the hip with expanded sacral ribs. The latter sacral rib is a single unit without a bifurcated structure. The tail is long, with at least 30 and possible up to 50 caudal vertebrae . The first few caudals are large, with closely interlinked zygapophyses and widely projecting pleurapophyses (a term for transverse processes lacking ribs). The length of
5730-475: A paper read at the Geological Society of London , during the same meeting in which for the first time a dinosaur was named, Megalosaurus . The two finds revealed the unique and bizarre build of the animals, in 1832 by Professor Buckland likened to "a sea serpent run through a turtle". In 1824, Conybeare also provided a specific name to Plesiosaurus : dolichodeirus , meaning "longneck". In 1848,
5921-415: A particularly large nasolacrimal duct , a tubular channel opening out of the rear of the lacrimal. The frontals are quite wide and form much of the upper edge of the orbit, a condition akin to T. longobardicus . However, the paired frontals meet along a straight suture with a low ridge on the lower (internal) surface, in contrast to T. longobardicus , where the frontals meet at an interdigitating suture with
Tanystropheus - Misplaced Pages Continue
6112-483: A plesiosaur skeleton, which he donated to Cope. Cope attempted to reconstruct the animal on the assumption that the longer extremity of the vertebral column was the tail, the shorter one the neck. He soon noticed that the skeleton taking shape under his hands had some very special qualities: the neck vertebrae had chevrons and with the tail vertebrae the joint surfaces were orientated back to front. Excited, Cope concluded to have discovered an entirely new group of reptiles:
6303-442: A prominent neural spine and robust zygapophyses, also unlike its predecessors. The 13th vertebra has long been assumed to be the first dorsal (torso) vertebra. This was justified by its general stout shape and supposedly dichocephalous (two-headed) rib facets, unlike the cervicals. However, specimen GMPKU-P-1527 has shown that the 13th vertebra's rib simply has a single wide articulation and an unconnected forward branch, more similar to
6494-429: A quantitative manner, by collecting a set of characteristics in sampled species and then using computational models to find the simplest ( most parsimonious ) path evolution could take to produce that character distribution. Cladistics stabilized and defined a fundamental split in the family tree of reptiles: one side of the family tree, Lepidosauromorpha , leads to lepidosaurs such as squamates (lizards and snakes) and
6685-655: A quarry at Fulbeck in Lincolnshire and had been used, with the fossil at its underside, to reinforce the slope of a watering-hole in Elston in Nottinghamshire . After the strange bones it contained had been discovered, it was displayed in the local vicarage as the remains of a sinner drowned in the Great Flood . Stukely affirmed its " diluvial " nature but understood it represented some sea creature, perhaps
6876-687: A rather small short-necked form. During the earliest Jurassic, the subgroup with the most species was the Rhomaleosauridae , a possibly very basal split-off of species which were also short-necked. Plesiosaurs in this period were at most five metres (sixteen feet) long. By the Toarcian , about 180 million years ago, other groups, among them the Plesiosauridae , became more numerous and some species developed longer necks, resulting in total body lengths of up to ten metres (33 feet). In
7067-407: A shallowly concave outer edge. Like the frontals, the paired parietals are seemingly separate bones, unfused to each other in every member of the species. A large hole, the pineal foramen (sometimes called the parietal foramen), is present at the midline of the skull between the front part of each parietal. When seen from below, a pair of curved crests along the frontals and parietals mark the edge of
7258-423: A single central narial opening. Unlike T. longobardicus , T. hydroides has a nearly vertical rear edge of the premaxilla, without a postnarial process. The maxilla is low, with a large and rectangular front portion. There is a perforation near the front of the bone, which would have been penetrated by the tenth dentary tooth when the mouth was closed. Towards the rear, the maxilla develops a concave edge overlooking
7449-507: A single row of 15 relatively large curved teeth along the outer edge of the bone, adjacent to the elongated choanae (internal openings of the nasal cavity ). Most other archosauromorphs, T. longobardicus included, have restricted vomers with rows of minuscule teeth. The rest of the palate is completely toothless in T. hydroides , even the palatines and pterygoids, which bear tooth rows in most early archosauromorphs. The pterygoids are also unusual for their broad palatal ramus (front plate) and
7640-433: A small fourth distal tarsal and a minuscule third distal tarsal. There are five closely appressed metatarsals (foot bones), with the fourth and third being the longest. Though the first four metatarsals are slender and similar in length, the fifth (outermost) is very stout and subtly hooked, slotting into the ankle along a smooth joint. The estimated phalangeal formula (joints per toe) is 2-3-4-5-4. The first phalange of
7831-428: A smaller and less specialized reptile found in the same geological strata. Beyond this conclusion, Peyer initially suggested that Tanystropheus was related to other long-necked Triassic reptiles. Sauropterygians such as plesiosaurs and nothosaurs were one possibility, and another was the fragmentary German reptile Trachelosaurus . Later, Peyer classified Tanystropheus and Macrocnemus closer to "protorosaurs",
Tanystropheus - Misplaced Pages Continue
8022-536: A specific species. The type species of Tanystropheus is T. conspicuus , a dubious name applied to particularly large fossils from Germany and Poland . Complete skeletons are common in the Besano Formation at Monte San Giorgio , on the border of Italy and Switzerland . Monte San Giorgio fossils belong to two species: the smaller T. longobardicus and the larger T. hydroides . These two species were formally differentiated in 2020 primarily on
8213-530: A term initially used for Permian reptiles such as Protorosaurus and Araeoscelis . In the early and mid-20th century, it was commonplace for Permian and Triassic reptiles of uncertain affinity to intermingle together in classification schemes. Names such as " Eosuchia ", " Euryapsida ", " Younginiformes ", " Protorosauria ", and others were all applied by different authors with little consistency. The Early Triassic reptile Prolacerta , from South Africa , also became involved upon its discovery. Prolacerta
8404-416: A three-lobed contact with the nasals. The sutures between the frontals and their neighboring bones are coarse and interdigitating (interlocking). A small triangular bone, the postfrontal , wedges behind the rear outer corner of each frontal. A pair of larger plate-like bones, the parietals , sit directly behind the frontals on the skull roof. In T. longobardicus , the parietals are fairly broad and flat, with
8595-406: A transverse ridge, similar to allokotosaurs and archosauriforms . The parabasisphenoid (front lower component) is less specialized; it lies flat and tapers forwards to a blade-like cultriform process. The rear part of the bone has a deep triangular excavation (known as a median pharyngeal recess) on its underside, flanked by low crests and a pair of small basipterygoid processes (knobs connecting to
8786-450: A while, Marsh suggested that a simpler explanation of the strange build would be that Cope had reversed the vertebral column relative to the body as a whole. When Cope reacted indignantly to this suggestion, Leidy silently took the skull and placed it against the presumed last tail vertebra, to which it fitted perfectly: it was in fact the first neck vertebra, with still a piece of the rear skull attached to it. Mortified, Cope tried to destroy
8977-408: A whole. The lower jaw is slender, and most of its length is devoted to the toothed dentary bone. The dentary is downturned at its tip and its outer surface is dotted with a row of prominent foramina (blood vessel pits). There are up to 19 teeth in the dentary. Most commonly, the first six teeth are prominent conical fangs, akin to the premaxilla, while the remainder are small and tricuspid, akin to
9168-920: Is a vertebra from the lower Carnian Fusea site in Friuli, Italy . In 2015, a large Tanystropheus cervical vertebra was described from the Economy Member of the Wolfville Formation , in the Bay of Fundy of Nova Scotia, Canada . The Wolfville Formation spans the Anisian to Carnian stages, and the Economy Member is likely Middle Triassic (Anisian-Ladinian) in age. It is a rare example of predominantly freshwater strata preserving Tanystropheus fossils. Tanystropheus -like tanystropheid fossils are known from another freshwater formation in North America:
9359-804: Is also found in Bulgaria near the Greek-Bulgarian border. Greek is also spoken worldwide by the sizable Greek diaspora which has notable communities in the United States , Australia , Canada , South Africa , Chile , Brazil , Argentina , Russia , Ukraine , the United Kingdom , and throughout the European Union , especially in Germany . Historically, significant Greek-speaking communities and regions were found throughout
9550-773: Is an Indo-European language, constituting an independent Hellenic branch within the Indo-European language family. It is native to Greece , Cyprus , Italy (in Calabria and Salento ), southern Albania , and other regions of the Balkans , Caucasus , the Black Sea coast, Asia Minor , and the Eastern Mediterranean . It has the longest documented history of any Indo-European language, spanning at least 3,400 years of written records. Its writing system
9741-479: Is an Indo-European language, but also includes a number of borrowings from the languages of the populations that inhabited Greece before the arrival of Proto-Greeks, some documented in Mycenaean texts ; they include a large number of Greek toponyms . The form and meaning of many words have changed. Loanwords (words of foreign origin) have entered the language, mainly from Latin, Venetian , and Turkish . During
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#17327802173789932-471: Is broader and flatter than that of T. longobardicus . The first five of six teeth in the premaxilla are very large and fang-like, forming an interlocking "fish trap" similar to Dinocephalosaurus and many sauropterygians such as plesiosaurs and nothosaurs . All teeth in the skull have a single cusp which is sharp, curved, and unserrated. They have an oval-shaped cross section and shallow subthecodont implantation. Like T. longobardicus , T. hydroides has
10123-408: Is called a " stem clade ". Such a definition has the advantage that it is easier to include all species with a certain morphology . Plesiosauria was in 2010 by Hillary Ketchum and Roger Benson defined as such a stem-based taxon : "all taxa more closely related to Plesiosaurus dolichodeirus and Pliosaurus brachydeirus than to Augustasaurus hagdorni ". Ketchum and Benson (2010) also coined
10314-464: Is currently on display at the geological museum of South Dakota School of Mines and Technology . In 2008, fossil remains of an undescribed plesiosaur that was named Predator X , now known as Pliosaurus funkei , were unearthed in Svalbard . It had a length of 12 m (39 ft), and its bite force of 149 kilonewtons (33,000 lb f ) is one of the most powerful known. In December 2017,
10505-403: Is floored and reinforced by a similar bone: the prearticular . In Tanystropheus species with known skull material, both the articular and prearticular contribute equally to a segment of the jaw extending back beyond the level of the jaw joint. This projection, known as a retroarticular process, is enlarged to a similar degree to that of early rhynchosaurs. The skull of Tanystropheus hydroides
10696-413: Is formed by the supraoccipitals , which were presumably fused together as a continuous surface sloping smoothly down to the foramen magnum. In the lower jaw, the dentaries meet each other at a robust symphysis with an interdigitating suture. The front end of the dentary hosts a prominent keel on its lower edge, a unique trait of the species. There are at least 18 dentary teeth; the first three are by far
10887-447: Is its hyperelongate neck, equivalent to the combined length of the body and tail. Tanystropheus has 13 cervical (neck) vertebrae , most of which are massive, though the two closest to the head are smaller and less strongly developed. The atlas (first cervical), which connects to the skull, is a small, four-part bone complex. It consists of an atlantal intercentrum (small lower component) and pleurocentrum (large lower component), and
11078-671: Is larger, with a small axial intercentrum followed by a much larger axial pleurocentrum. The axial pleurocentrum is longer than tall, has a low neural spine set forwards, and small prezygapophyses (front articular plates). The large postzygophyses (rear articular plates) are separated by a broad trough and support pointed epipophyses (additional projections). The third to eleventh cervicals are hyperelongate in T. longobardicus and T. hydroides , ranging from three to 15 times longer than tall. They are somewhat less elongated in T. antiquus , less than 6 times longer than tall. The cervicals gradually increase in size and proportional length, with
11269-508: Is low and extends to a tapered point at the rear. The pubis (lower front hip blade) is vertically oriented, with a small but distinct obturator foramen and a concave rear edge. The lower front tip of the large, fan-shaped ischium (lower rear hip blade) converges towards the pubis, but does not contact it. The large oval-shaped gap between the pubis and ischium is known as the thyroid foramen . Two pairs of large, curved bones, known as heterotopic ossifications or postcloacal bones, sit behind
11460-469: Is nestled further down along the maxilla. The lower edge of the orbit is formed by the jugal , a bone with a slender anterior process (front branch) and a somewhat broader dorsal process (upper branch). There is also a very short pointed posterior process (rear branch) which ends freely and fails to contact any other bone. The shape of the jugal in Tanystropheus is typical for early archosauromorphs;
11651-687: Is now preserved by the British Geological Survey as specimen BGS GSM 26035. The same year, commercial fossil collector Mary Anning and her family uncovered an almost complete skeleton at Lyme Regis in Dorset , England, on what is today called the Jurassic Coast . It was acquired by the Duke of Buckingham , who made it available to the geologist William Buckland . He in turn let it be described by Conybeare on 20 February 1824 in
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#173278021737811842-600: Is one of the most well-described non- archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae , a clade collecting many long-necked Triassic archosauromorphs previously described as " protorosaurs " or " prolacertiforms ". Tanystropheus contains at least two valid species as well as fossils which cannot be referred to
12033-412: Is positioned near the straight contact with the frontals, one of the few similarities with T. longobardicus . Strong supratemporal fossae excavate into the outer edge of the parietal and define a low sagittal crest along the midline of the skull. This trend is shared with other large archosauromorphs, like Dinocephalosaurus and Azendohsaurus . The supratemporal fenestrae (upper skull holes behind
12224-519: Is protected and promoted officially as a regional and minority language in Armenia, Hungary , Romania, and Ukraine. It is recognized as a minority language and protected in Turkey by the 1923 Treaty of Lausanne . The phonology , morphology , syntax , and vocabulary of the language show both conservative and innovative tendencies across the entire attestation of the language from the ancient to
12415-428: Is rather stiff and ungainly, suggesting a reliance on water. Conversely, the limbs and tail lack most adaptations for swimming and closely resemble their equivalents in terrestrial reptiles. Recent studies have supported an intermediate position, reconstructing Tanystropheus as an animal equally capable on land and in the water. Despite its length, the neck was lightweight and stabilized by tendons , so it would not been
12606-448: Is seen in few other archosauromorphs , namely rhynchosaurs , most allokotosaurs , modern crocodilians , and Teyujagua . The maxilla is triangular, reaching its maximum height at mid-length and tapering to the front and rear. There are up to 14 or 15 teeth in the maxilla, though some individuals have fewer. T. longobardicus is a reptile with heterodont dentition, meaning that it had more than one type of tooth shape. In contrast to
12797-442: Is similar in shape to the humerus and lacks a distinct olecranon (elbow projection). There are four carpals (wrist bones): the ulnare , radiale , and two distal carpals. The ulnare and radiale are large and cuboid, enclosing a small foramen (gap) between them. The larger outer distal carpal connects to metacarpals III and IV, while the much smaller inner distal carpal connects to metacarpals II and III. Metacarpals III and IV are
12988-947: Is sometimes called aljamiado , as when Romance languages are written in the Arabic alphabet. Article 1 of the Universal Declaration of Human Rights in Greek: Transcription of the example text into Latin alphabet : Article 1 of the Universal Declaration of Human Rights in English: Proto-Greek Mycenaean Ancient Koine Medieval Modern Plesiosaur The Plesiosauria or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles , belonging to
13179-526: Is spoken by at least 13.5 million people today in Greece, Cyprus, Italy, Albania, Turkey , and the many other countries of the Greek diaspora . Greek roots have been widely used for centuries and continue to be widely used to coin new words in other languages; Greek and Latin are the predominant sources of international scientific vocabulary . Greek has been spoken in the Balkan peninsula since around
13370-763: Is still used internationally for the writing of Ancient Greek . In Greek, the question mark is written as the English semicolon, while the functions of the colon and semicolon are performed by a raised point (•), known as the ano teleia ( άνω τελεία ). In Greek the comma also functions as a silent letter in a handful of Greek words, principally distinguishing ό,τι ( ó,ti , 'whatever') from ότι ( óti , 'that'). Ancient Greek texts often used scriptio continua ('continuous writing'), which means that ancient authors and scribes would write word after word with no spaces or punctuation between words to differentiate or mark boundaries. Boustrophedon , or bi-directional text,
13561-650: Is the Greek alphabet , which has been used for approximately 2,800 years; previously, Greek was recorded in writing systems such as Linear B and the Cypriot syllabary . The alphabet arose from the Phoenician script and was in turn the basis of the Latin , Cyrillic , Coptic , Gothic , and many other writing systems. The Greek language holds a very important place in the history of the Western world. Beginning with
13752-495: The Eastern Mediterranean , in what are today Southern Italy , Turkey , Cyprus , Syria , Lebanon , Israel , Palestine , Egypt , and Libya ; in the area of the Black Sea , in what are today Turkey, Bulgaria , Romania , Ukraine , Russia , Georgia , Armenia , and Azerbaijan ; and, to a lesser extent, in the Western Mediterranean in and around colonies such as Massalia , Monoikos , and Mainake . It
13943-516: The Indo-Iranian languages (see Graeco-Aryan ), but little definitive evidence has been found. In addition, Albanian has also been considered somewhat related to Greek and Armenian, and it has been proposed that they all form a higher-order subgroup along with other extinct languages of the ancient Balkans; this higher-order subgroup is usually termed Palaeo-Balkan , and Greek has a central position in it. Linear B , attested as early as
14134-691: The Middle Triassic epoch ( Anisian and Ladinian stages), with some exceptions. For example, a vertebra from Nova Scotia was recovered from primarily freshwater sediments. The youngest fossils in the genus are a pair of well-preserved skeletons from the Zhuganpo Formation , a geological unit in China which dates to the earliest part of the Late Triassic (early Carnian stage). The oldest putative fossils belong to "T. antiquus" ,
14325-515: The Middle Triassic , from the latest Anisian to middle Ladinian stages. Though the fossils were initially given the name Macroscelosaurus by Count Georg Zu Münster , the publication containing this name is lost and its genus is considered a nomen oblitum . In 1855, Hermann von Meyer supplied the name Tanystropheus conspicuus , the type species of Tanystropheus , to the fossils. They were later regarded as Tanystropheus fossils undiagnostic relative to other species, rendering T. conspicuus
14516-650: The Polycotylidae , which had short necks and peculiarly elongated heads with narrow snouts. During the Late Cretaceous, the elasmosaurids still had many species. All plesiosaurs became extinct as a result of the K-T event at the end of the Cretaceous period, approximately 66 million years ago. In modern phylogeny , clades are defined groups that contain all species belonging to a certain branch of
14707-655: The Sauria , particularly the crocodile, than Ichthyosaurus , which had the form of a more lowly fish. The name should thus be rather read as "approaching the Sauria" or "near reptile" than as "near lizard". Parts of the specimen are still present in the Oxford University Museum of Natural History . Soon afterwards, the morphology became much better known. In 1823, Thomas Clark reported an almost complete skull, probably belonging to Thalassiodracon , which
14898-818: The Sauropterygia . Plesiosaurs first appeared in the latest Triassic Period , possibly in the Rhaetian stage, about 203 million years ago. They became especially common during the Jurassic Period, thriving until their disappearance due to the Cretaceous–Paleogene extinction event at the end of the Cretaceous Period, about 66 million years ago. They had a worldwide oceanic distribution, and some species at least partly inhabited freshwater environments. Plesiosaurs were among
15089-477: The Streptosauria or "Turned Saurians", which would be distinguished by reversed vertebrae and a lack of hindlimbs, the tail providing the main propulsion. After having published a description of this animal, followed by an illustration in a textbook about reptiles and amphibians, Cope invited Marsh and Joseph Leidy to admire his new Elasmosaurus platyurus . Having listened to Cope's interpretation for
15280-636: The Upper Triassic . One of these, the Nothosauroidea , kept functional elbow and knee joints; but the other, the Pistosauria , became more fully adapted to a sea-dwelling lifestyle. Their vertebral column became stiffer and the main propulsion while swimming no longer came from the tail but from the limbs, which changed into flippers. The Pistosauria became warm-blooded and viviparous , giving birth to live young. Early, basal , members of
15471-535: The Villány Mountains of Hungary . The most well-preserved Tanystropheus fossils outside of Monte San Giorgio come from the Guizhou province of China , as described by Li (2007) and Rieppel (2010). They are also among the youngest and easternmost fossils in the genus, hailing from the upper Ladinian or lower Carnian Zhuganpo Formation . Although the postcrania is complete and indistinguishable from
15662-402: The coracoid (lower shoulder blade), which is a large oval-shaped plate with a broad glenoid facet (shoulder socket). The humerus (upper arm bone) is straight and slightly constricted at the middle. Near the elbow it is expanded and twisted, with an ectepicondylar groove on its outer edge. The radius (outer forearm bone) is slender and somewhat curved, while the ulna (inner forearm bone)
15853-414: The forebrain , as defined by a bulbous central hollow. The eye was supported by more than 10 rectangular ossicles (tiny plate-like bones) connecting into a scleral ring , though a full reconstruction of the ring, with 18 ossicles, is conjectural. Few details of the braincase and palate (bony roof of the mouth) are known for T. longobardicus . The scant available evidence suggests that these regions of
16044-492: The nominal and verbal systems. The major change in the nominal morphology since the classical stage was the disuse of the dative case (its functions being largely taken over by the genitive ). The verbal system has lost the infinitive , the synthetically -formed future, and perfect tenses and the optative mood . Many have been replaced by periphrastic ( analytical ) forms. Pronouns show distinctions in person (1st, 2nd, and 3rd), number (singular, dual , and plural in
16235-402: The phylogeny , the evolutionary relationships, and the morphology , the way the animal is built, long-necked forms are therefore called "plesiosauromorph" and short-necked forms are called "pliosauromorph", without the "plesiosauromorph" species necessarily being more closely related to each other than to the "pliosauromorph" forms. The latest common ancestor of the Plesiosauria was probably
16426-837: The sister taxon to Tanytrachelos , a much smaller tanystropheid from Virginia . Another small tanystropheid, Cosesaurus from Spain , is allied with the Tanystropheus + Tanytrachelos clade in many analyses of the 1980s and 1990s. Within Archosauromorpha, "prolacertiforms" are joined by several other groups. The clade Archosauriformes is a diverse archosauromorph subset including crown group archosaurs and their predatory close relatives such as Euparkeria and Proterosuchus . Stocky Triassic herbivores like rhynchosaurs , Trilophosaurus , and azendohsaurids additionally qualify as archosauromorphs. The bizarre chameleon -like drepanosaurs were also included by many analyses, though more recently they have been reinterpreted as
16617-585: The tuatara . The other side, Archosauromorpha , leads to archosaurs. Cladistics was one of many lines of evidence that helped to demonstrate the dinosaurian origin of birds. This left crocodilians and birds as the two surviving archosaur groups. A series of phylogenetic analyses in the late 1980s and 1990s strongly supported the proposal of Gow (1975). Tanystropheus , Macrocnemus , Protorosaurus , and Prolacerta were always placed as members of Archosauromorpha, closer to archosaurs than to squamates. "Protorosauria" and "Prolacertiformes" were used interchangeably for
16808-401: The "plesiosauromorph" build, had (sometimes extremely) long necks and small heads; these were relatively slow and caught small sea animals. Other species, some of them reaching a length of up to seventeen metres, had the "pliosauromorph" build with a short neck and a large head; these were apex predators , fast hunters of large prey. The two types are related to the traditional strict division of
16999-412: The 1970s, not long after the field of paleontology was reinvigorated by the " dinosaur renaissance " in the 1960s and beyond. In the 1980s, the advent of cladistics saw a paradigm shift in the field of taxonomy , emphasizing monophyletic clades (all-encompassing groups defined by shared ancestry) over other categorization styles. Phylogenetic analyses were invented to evaluate reptile evolution in
17190-573: The 1990s near Herschel, Saskatchewan were found to be a new species, by Dr. Tamaki Sato, a Japanese vertebrate paleontologist. In 2006, a combined team of American and Argentinian investigators (the latter from the Argentinian Antarctic Institute and the La Plata Museum ) found the skeleton of a juvenile plesiosaur measuring 1.5 metres (4 ft 11 in) in length on Vega Island in Antarctica. The fossil
17381-470: The 3rd millennium BC, or possibly earlier. The earliest written evidence is a Linear B clay tablet found in Messenia that dates to between 1450 and 1350 BC, making Greek the world's oldest recorded living language . Among the Indo-European languages, its date of earliest written attestation is matched only by the now-extinct Anatolian languages . The Greek language is conventionally divided into
17572-690: The Anisian-age Moenkopi Formation of Arizona and New Mexico. Several new tanystropheid genera have been named from former Tanystropheus fossils. Fossils from the Anisian Röt Formation in Germany, previously referred to Tanystropheus antiquus, were named as a new genus and species in 2006: Amotosaurus rotfeldensis . In 2011, fossils from the Lipovskaya Formation of Russia were given
17763-475: The British Museum of Natural History; this is today referred to Thalassiodracon . In the early nineteenth century, plesiosaurs were still poorly known and their special build was not understood. No systematic distinction was made with ichthyosaurs , so the fossils of one group were sometimes combined with those of the other to obtain a more complete specimen. In 1821, a partial skeleton discovered in
17954-486: The Greek alphabet since approximately the 9th century BC. It was created by modifying the Phoenician alphabet , with the innovation of adopting certain letters to represent the vowels. The variant of the alphabet in use today is essentially the late Ionic variant, introduced for writing classical Attic in 403 BC. In classical Greek, as in classical Latin, only upper-case letters existed. The lower-case Greek letters were developed much later by medieval scribes to permit
18145-425: The Greek language are often emphasized. Although Greek has undergone morphological and phonological changes comparable to those seen in other languages, never since classical antiquity has its cultural, literary, and orthographic tradition been interrupted to the extent that one can speak of a new language emerging. Greek speakers today still tend to regard literary works of ancient Greek as part of their own rather than
18336-495: The Greek language was the Cypriot syllabary (also a descendant of Linear A via the intermediate Cypro-Minoan syllabary ), which is closely related to Linear B but uses somewhat different syllabic conventions to represent phoneme sequences. The Cypriot syllabary is attested in Cyprus from the 11th century BC until its gradual abandonment in the late Classical period, in favor of the standard Greek alphabet. Greek has been written in
18527-629: The Greek verb have likewise remained largely the same over the course of the language's history but with significant changes in the number of distinctions within each category and their morphological expression. Greek verbs have synthetic inflectional forms for: Many aspects of the syntax of Greek have remained constant: verbs agree with their subject only, the use of the surviving cases is largely intact (nominative for subjects and predicates, accusative for objects of most verbs and many prepositions, genitive for possessors), articles precede nouns, adpositions are largely prepositional, relative clauses follow
18718-685: The Greek-Albanian border. A significant percentage of Albania's population has knowledge of the Greek language due in part to the Albanian wave of immigration to Greece in the 1980s and '90s and the Greek community in the country. Prior to the Greco-Turkish War and the resulting population exchange in 1923 a very large population of Greek-speakers also existed in Turkey , though very few remain today. A small Greek-speaking community
18909-491: The Museo Cantonale di Scienze Naturali di Lugano (MCSN). Rupert Wild reviewed and redescribed all specimens known at the time via several large monographs in 1973/4 and 1980. In 2005, Silvio Renesto described a T. longobardicus specimen from Switzerland which preserved the impressions of skin and other soft tissue. Five new MSNM specimens of T. longobardicus were described by Stefania Nosotti in 2007, allowing for
19100-425: The Plesiosauria into two suborders, the long-necked Plesiosauroidea and the short-neck Pliosauroidea . Modern research, however, indicates that several "long-necked" groups might have had some short-necked members or vice versa. Therefore, the purely descriptive terms "plesiosauromorph" and "pliosauromorph" have been introduced, which do not imply a direct relationship. "Plesiosauroidea" and "Pliosauroidea" today have
19291-404: The acute during the late 20th century, and it has only been retained in typography . After the writing reform of 1982, most diacritics are no longer used. Since then, Greek has been written mostly in the simplified monotonic orthography (or monotonic system), which employs only the acute accent and the diaeresis. The traditional system, now called the polytonic orthography (or polytonic system),
19482-402: The ancient language; singular and plural alone in later stages), and gender (masculine, feminine, and neuter), and decline for case (from six cases in the earliest forms attested to four in the modern language). Nouns, articles, and adjectives show all the distinctions except for a person. Both attributive and predicative adjectives agree with the noun. The inflectional categories of
19673-414: The archosauromorph subgroup encompassing these superficially lizard-like reptiles. Some authors preferred "Protorosauria" for its priority . Most others used "Prolacertiformes" arguing that "Protorosauria" was a name that carried too much historical baggage, since it had previously encompassed non-archosauromorph "euryapsids" like Araeoscelis . As a "prolacertiform", Tanystropheus is typically considered
19864-458: The basis of their strongly divergent skull anatomy. When T. longobardicus was first described in 1886, it was initially mistaken for a pterosaur and given the name " Tribelesodon ". Starting in the 1920s, systematic excavations at Monte San Giorgio unearthed many more Tanystropheus fossils, revealing that the putative wing bones of "Tribelesodon" were actually neck vertebrae. Most Tanystropheus fossils hail from marine or coastal deposits of
20055-532: The cervical ribs than the dorsal ribs. The elongation of Tanystropheus 's neck is mostly a consequence of particular vertebrae lengthening. This is a contrast to trachelosaurids such as Dinocephalosaurus , which achieve a long neck by the addition of numerous cervicals, for a total cervical count exceeding 30. Nevertheless, Tanystropheus does have more vertebrae in its neck than typical archosauromorphs. Protorosaurus , for example, has only seven cervicals, while Macrocnemus and Prolacerta have eight. To achieve
20246-410: The cervicals. Though their neural spines are taller than those of the cervicals, they are still rather short. The dorsal ribs are double-headed close to the shoulder and single-headed in the rest of the torso, sitting on stout transverse processes projecting outwards from the front half of each vertebra. More than 20 angled rows of gastralia extend along the belly, each gastral element represented by
20437-537: The collection of Colonel Thomas James Birch , was described by William Conybeare and Henry Thomas De la Beche , and recognised as representing a distinctive group. A new genus was named, Plesiosaurus . The generic name was derived from the Greek πλήσιος, plèsios , "closer to" and the Latinised saurus , in the meaning of "saurian", to express that Plesiosaurus was in the Chain of Being more closely positioned to
20628-479: The collector Thomas Hawkins : Memoirs of Ichthyosauri and Plesiosauri of 1834 and The Book of the Great Sea-Dragons of 1840. Hawkins entertained a very idiosyncratic view of the animals, seeing them as monstrous creations of the devil, during a pre-Adamitic phase of history. Hawkins eventually sold his valuable and attractively restored specimens to the British Museum of Natural History. During
20819-532: The earlier Plesiosaurus dolichodeirus as it is derived from πλεῖος, pleios , "more fully", reflecting that according to Owen it was closer to the Sauria than Plesiosaurus . Its specific name means "with a short neck". Later, the Pliosauridae were recognised as having a morphology fundamentally different from the plesiosaurids. The family Plesiosauridae had already been coined by John Edward Gray in 1825. In 1835, Henri Marie Ducrotay de Blainville named
21010-404: The early 19th century that was used for literary and official purposes in the newly formed Greek state. In 1976, Dimotiki was declared the official language of Greece, after having incorporated features of Katharevousa and thus giving birth to Standard Modern Greek , used today for all official purposes and in education . The historical unity and continuing identity between the various stages of
21201-460: The early twenty-first century, with about three or four plesiosaurs being named each year. This implies that about half of the known plesiosaurs are relatively new to science, a result of a far more intense field research. Some of this is taking place away from the traditional areas, e.g. in new sites developed in New Zealand , Argentina , Chile , Norway , Japan , China and Morocco , but
21392-440: The edge of the cranium. In T. longobardicus , the paroccipital processes are shorter and narrower at their base. The stapes , a bone which transmits vibrations from the ear to the braincase, is slender and splits into two small prongs where it contacts the opisthotic. The opisthotic merges forwards into the prootic , which extensively contacts the parabasisphenoid and hosts a range of larger nerve foramina. The prootic forms much of
21583-579: The entire edition of the textbook and, when this failed, immediately published an improved edition with a correct illustration but an identical date of publication. He excused his mistake by claiming that he had been misled by Leidy himself, who, describing a specimen of Cimoliasaurus , had also reversed the vertebral column. Marsh later claimed that the affair was the cause of his rivalry with Cope: "he has since been my bitter enemy". Both Cope and Marsh in their rivalry named many plesiosaur genera and species, most of which are today considered invalid. Around
21774-618: The epics of Homer , ancient Greek literature includes many works of lasting importance in the European canon . Greek is also the language in which many of the foundational texts in science and philosophy were originally composed. The New Testament of the Christian Bible was also originally written in Greek. Together with the Latin texts and traditions of the Roman world , the Greek texts and Greek societies of antiquity constitute
21965-463: The evolutionary tree . One way to define a clade is to let it consist of the last common ancestor of two such species and all its descendants. Such a clade is called a " node clade ". In 2008, Patrick Druckenmiller and Anthony Russell in this way defined Plesiosauria as the group consisting of the last common ancestor of Plesiosaurus dolichocheirus and Peloneustes philarchus and all its descendants. Plesiosaurus and Peloneustes represented
22156-428: The eye) are wide and semi-triangular, exposed almost entirely from above. The postorbital has large and blocky ventral and medial processes (lower and inward branches), which meet at a sharper angle than in any other early archosauromorph. The jugal, conversely, is basically indistinguishable from that of T. longobardicus . The squamosal is deep and rectangular when viewed from the side, with little differentiation between
22347-419: The fifth toe was very long, filling a metatarsal-like role as seen in other tanystropheids. Knowledge on the anatomy of Tanystropheus was transformed by Bernhard Peyer 's discoveries in the 1920s and 1930s, but its relationship to other reptiles remained enigmatic for much of the 20th century. Most paleontologists (including modern authorities) agree that Tanystropheus was closely related to Macrocnemus ,
22538-511: The first fossil reptiles discovered. In the beginning of the nineteenth century, scientists realised how distinctive their build was and they were named as a separate order in 1835. The first plesiosaurian genus, the eponymous Plesiosaurus , was named in 1821. Since then, more than a hundred valid species have been described. In the early twenty-first century, the number of discoveries has increased, leading to an improved understanding of their anatomy, relationships and way of life. Plesiosaurs had
22729-573: The first fossils of extinct reptiles recognised as such. In 1605, Richard Verstegen of Antwerp illustrated in his A Restitution of Decayed Intelligence plesiosaur vertebrae that he referred to fishes and saw as proof that Great Britain was once connected to the European continent. The Welshman Edward Lhuyd in his Lithophylacii Brittannici Ichnographia from 1699 also included depictions of plesiosaur vertebrae that again were considered fish vertebrae or Ichthyospondyli . Other naturalists during
22920-419: The first half of the nineteenth century, the number of plesiosaur finds steadily increased, especially through discoveries in the sea cliffs of Lyme Regis. Sir Richard Owen alone named nearly a hundred new species. The majority of their descriptions were, however, based on isolated bones, without sufficient diagnosis to be able to distinguish them from the other species that had previously been described. Many of
23111-452: The following periods: In the modern era, the Greek language entered a state of diglossia : the coexistence of vernacular and archaizing written forms of the language. What came to be known as the Greek language question was a polarization between two competing varieties of Modern Greek: Dimotiki , the vernacular form of Modern Greek proper, and Katharevousa , meaning 'purified', a compromise between Dimotiki and Ancient Greek developed in
23302-421: The forelimbs, though similar in overall structure and proportions. The femur (thigh bone) is long, slender, and sigmoid (curved at both ends). It has a longitudinal muscle crest for muscle attachment (the internal trochanter ) on its underside, and it contacts the acetabulum (hip socket) at a broad smooth joint. The tibia and fibula (shin bones) are straight, with the former much thicker and more expanded at
23493-638: The fossils of Monte San Giorgio, no skull material is preserved, and their younger age precludes unambiguous placement into any Tanystropheus species. The Chinese material includes a large morphotype ( T. hydroides? ) specimen, GMPKU-P-1527, and an indeterminate juvenile skeleton, IVPP V 14472. Indeterminate Tanystropheus remains are also known from the Jilh Formation of Saudi Arabia and various Anisian-Ladinian sites in Spain , France , Italy, and Switzerland. The youngest Tanystropheus fossil in Europe
23684-425: The front edge of the interclavicle , a plate-like bone at the center of the chest with a rhombic (broad, diamond-shaped) front region followed by a long stalk at the rear. The interclavicle is rarely preserved and its connections to the rest of the pectoral girdle are mostly inferred from Macrocnemus . The scapula (upper shoulder blade) has the form of a large semicircular plate on a short, broad stalk. It lies above
23875-404: The front edge of the paroccipital process, akin to the condition in archosauriforms. Another archosauriform-like feature is the presence of a laterosphenoid , an additional braincase component in front of the prootic and above the exit hole for the trigeminal nerve (also known as cranial nerve V). The laterosphenoid is small, similar to that of Azendohsaurus . The upper rear part of the braincase
24066-587: The fully aquatic plesiosaurs . 19th century excavations at Monte San Giorgio , a UNESCO world heritage site on the Italy - Switzerland border, revealed a fragmentary fossil of an animal with three-cusped (tricuspid) teeth and elongated bones. Monte San Giorgio preserves the Besano Formation (also known as the Grenzbitumenzone), a late Anisian -early Ladinian lagerstätte recognised for its spectacular fossils. In 1886, Francesco Bassani interpreted
24257-423: The group, traditionally called " pistosaurids ", were still largely coastal animals. Their shoulder girdles remained weak, their pelves could not support the power of a strong swimming stroke, and their flippers were blunt. Later, a more advanced pistosaurian group split off: the Plesiosauria. These had reinforced shoulder girdles, flatter pelves, and more pointed flippers. Other adaptations allowing them to colonise
24448-470: The hips in about half of known specimens preserving the area. They occupy the base of the tail, a region which lacks chevrons. These bones are possibly sexually dimorphic , and have also been reported in the small American tanystropheid Tanytrachelos . Heterotopic ossifications may be linked to reproductive biology, supporting reproductive organs (if they belong to males) or an egg pouch (if they belong to females). The hindlimbs are significantly larger than
24639-405: The hypothesis that they represent separate species. A 2020 study found numerous differences between the skulls of large and small specimens, formalizing the proposal to divide the two into separate species. Moreover, a histological investigation revealed that one small specimen, PIMUZ T 1277, was a skeletally mature adult at a length of only 1.5 meters (4.9 ft). The larger one-cusped morphotype
24830-439: The infinitive entirely (employing a raft of new periphrastic constructions instead) and uses participles more restrictively. The loss of the dative led to a rise of prepositional indirect objects (and the use of the genitive to directly mark these as well). Ancient Greek tended to be verb-final, but neutral word order in the modern language is VSO or SVO. Modern Greek inherits most of its vocabulary from Ancient Greek, which in turn
25021-423: The jaw. Although it is plausible that a small coronoid bone could be present in front of the surangular, evidence is ambiguous at best for all Tanystropheus species. A sheathe-like bone, the angular , is well-exposed under the dentary and surangular, though sutures between these bones are difficult to interpret with certainty. The joint at the back of the jaw lies on the articular , a lumpy rectangular bone which
25212-426: The knee. The large proximal tarsals (ankle or heel bones contacting the shin) consist of a rounded calcaneum and a blocky astragalus , which meet each other along a straight or shallowly indented contact in most specimens. Like most non-aquatic reptiles, a set of small pebble-shaped distal tarsals are present between the proximal tarsals and the foot bones. Tanystropheus has a reduced number of distal tarsals: only
25403-403: The largest bones in the hand, followed closely by metacarpal II. Metacarpals I and V are both short. The hand's phalangeal formula (joints per finger) is 2-3-4-4-3. The terminal phalanges (fingertips) may have formed thick, blunt claws. The components of the pelvis (hip) are proportionally small, though their shape is unremarkable relative to other tanystropheids. The ilium (upper hip blade)
25594-402: The largest teeth in the skull, forming the lower half of the interlocking "fish trap" with the premaxilla. Most other teeth in the dentary are small, with the exception of the tenth tooth, which juts up to pierce the maxilla. The remainder of the jaw contains the same set of bones as in T. longobardicus , but some details differ in T. hydroides . For example, the splenial is plate-like and covers
25785-467: The late 15th century BC, was the first script used to write Greek. It is basically a syllabary , which was finally deciphered by Michael Ventris and John Chadwick in the 1950s (its precursor, Linear A , has not been deciphered and most likely encodes a non-Greek language). The language of the Linear B texts, Mycenaean Greek , is the earliest known form of Greek. Another similar system used to write
25976-484: The late 2010s. The proportions of T. antiquus fossils are easily distinguishable, and it is currently considered a valid species of archosauromorph, though its referral to the genus Tanystropheus has been questioned . The Gogolin Formation ranges from the upper Olenekian (latest part of the Early Triassic ) to the lower Anisian in age. Assuming they belong within Tanystropheus , the fossils of T. antiquus may be
26167-493: The locations of the more original discoveries have proven to be still productive, with important new finds in England and Germany. Some of the new genera are a renaming of already known species, which were deemed sufficiently different to warrant a separate genus name. In 2002, the " Monster of Aramberri " was announced to the press. Discovered in 1982 at the village of Aramberri , in the northern Mexican state of Nuevo León , it
26358-528: The longest plesiosaurs, reaching up to fifteen metres (fifty feet) in length due to very long necks containing as many as 76 vertebrae, more than any other known vertebrate. Pliosauridae were still present as is shown by large predators, such as Kronosaurus . At the beginning of the Late Cretaceous , the Ichthyosauria became extinct; perhaps a plesiosaur group evolved to fill their niches:
26549-533: The main subgroups of the Plesiosauroidea and the Pliosauroidea and were chosen for historical reasons; any other species from these groups would have sufficed. Another way to define a clade is to let it consist of all species more closely related to a certain species that one in any case wishes to include in the clade than to another species that one to the contrary desires to exclude. Such a clade
26740-413: The maxilla. There is some variation in the number of each tooth shape, and some individuals may have up to 11 conical teeth. The inner surface of the dentary is joined by a splint-shaped bone, the splenial , at its lower edge. The splenial was most likely not visible in lateral view. At its rear, the dentary seems to be partially overlapped by the surangular , a bone which comprises much of the rear part of
26931-640: The membership of Greece and Cyprus in the European Union, Greek is one of the organization's 24 official languages . Greek is recognized as a minority language in Albania, and used co-officially in some of its municipalities, in the districts of Gjirokastër and Sarandë . It is also an official minority language in the regions of Apulia and Calabria in Italy. In the framework of the European Charter for Regional or Minority Languages , Greek
27122-456: The middle Ladinian Erfurt Formation (Lettenkeuper) of Germany was described in 1846 as one of several fossils gathered under the name " Zanclodon laevis ". Though likely the first Tanystropheus fossil to be discovered, the vertebra is now lost, and surviving jaw fragments and other fossil scraps of " Zanclodon laevis" represent indeterminate archosauriforms with no relation to Tanystropheus . Tanystropheus vertebrae have also been found in
27313-468: The middle of the Jurassic, very large Pliosauridae evolved. These were characterized by a large head and a short neck, such as Liopleurodon and Simolestes . These forms had skulls up to three metres (ten feet) long and reached a length of up to seventeen metres (56 feet) and a weight of ten tonnes. The pliosaurids had large, conical teeth and were the dominant marine carnivores of their time. During
27504-482: The middle of the twentieth century, the USA remained an important centre of research, mainly through the discoveries of Samuel Paul Welles . Whereas during the nineteenth and most of the twentieth century, new plesiosaurs were described at a rate of three or four novel genera each decade, the pace suddenly picked up in the 1990s, with seventeen plesiosaurs being discovered in this period. The tempo of discovery accelerated in
27695-441: The modern period. The division into conventional periods is, as with all such periodizations, relatively arbitrary, especially because, in all periods, Ancient Greek has enjoyed high prestige, and the literate borrowed heavily from it. Across its history, the syllabic structure of Greek has varied little: Greek shows a mixed syllable structure, permitting complex syllabic onsets but very restricted codas. It has only oral vowels and
27886-504: The morphology of the genus. The Miedary locality represents an isolated brackish body of water close to the coast, and the abundance of Tanystropheus fossils suggests that it was an animal well-suited for this kind of habitat. In the late 1900s, Friedrich von Huene named several dubious Tanystropheus species from Germany and Poland. T. posthumus , from the Norian of Germany, was later reevaluated as an indeterminate theropod vertebra and
28077-593: The new genus Coelophysis . Authentic Tanystropheus specimens from the Makhtesh Ramon in Israel were described as a new species, T. haasi , in 2001. However, this species may be dubious due to the difficulty of distinguishing its vertebrae from T. conspicuus or T. longobardicus . Another new species, T. biharicus , was described from Romania in 1975. It has also been considered possibly synonymous with T. longobardicus . A Tanystropheus -like vertebra from
28268-632: The new genus and species Augustaburiania vatagini by A.G. Sennikov. He also named the new genus Protanystropheus for T. antiquus, though a few studies continued to retain that species within Tanystropheus . Tanystropheus fossai, from the Norian -age Argillite di Riva di Solto in Italy, was given its own genus Sclerostropheus in 2019. Tanystropheus was one of the longest known non-archosauriform archosauromorphs. Vertebrae referred to " T. conspicuus " may correspond to an animal up to five or six meters (16.4 to 20 feet ) in length . T. hydroides
28459-419: The new species described at this time have subsequently been invalidated . The genus Plesiosaurus is particularly problematic, as the majority of the new species were placed in it so that it became a wastebasket taxon . Gradually, other genera were named. Hawkins had already created new genera, though these are no longer seen as valid. In 1841, Owen named Pliosaurus brachydeirus . Its etymology referred to
28650-549: The ninth cervical typically being the largest vertebra in the skeleton. In general structure, the elongated cervicals resemble the axial pleurocentrum. However, the axis also has a keel on its underside and an incomplete neural canal, unlike its immediate successors. In the rest of the cervicals, all but the front of each neural spine is so low that it is barely noticeable as a thin ridge. The zygapophyses are closely set and tightly connected between vertebrae. The epipophyses develop into hooked spurs. The cervicals are also compressed from
28841-399: The noun they modify and relative pronouns are clause-initial. However, the morphological changes also have their counterparts in the syntax, and there are also significant differences between the syntax of the ancient and that of the modern form of the language . Ancient Greek made great use of participial constructions and of constructions involving the infinitive, and the modern variety lacks
29032-536: The objects of study of the discipline of Classics . During antiquity , Greek was by far the most widely spoken lingua franca in the Mediterranean world . It eventually became the official language of the Byzantine Empire and developed into Medieval Greek . In its modern form , Greek is the official language of Greece and Cyprus and one of the 24 official languages of the European Union . It
29223-706: The older periods of Greek, loanwords into Greek acquired Greek inflections, thus leaving only a foreign root word. Modern borrowings (from the 20th century on), especially from French and English, are typically not inflected; other modern borrowings are derived from Albanian , South Slavic ( Macedonian / Bulgarian ) and Eastern Romance languages ( Aromanian and Megleno-Romanian ). Greek words have been widely borrowed into other languages, including English. Example words include: mathematics , physics , astronomy , democracy , philosophy , athletics , theatre, rhetoric , baptism , evangelist , etc. Moreover, Greek words and word elements continue to be productive as
29414-681: The oldest in the genus. Specimens likely referable to T. antiquus are also known from throughout Germany and the fossiliferous Winterswijk site in the Netherlands . In the 1880s, E.D. Cope named three supposed new Tanystropheus species ( T. bauri , T. willistoni , and T. longicollis ) from the Late Triassic Chinle Formation in New Mexico . However, these fossils were later determined to be tail vertebrae belonging to theropod dinosaurs, which were named under
29605-466: The open seas included stiff limb joints; an increase in the number of phalanges of the hand and foot; a tighter lateral connection of the finger and toe phalanx series, and a shortened tail. From the earliest Jurassic , the Hettangian stage, a rich radiation of plesiosaurs is known, implying that the group must already have diversified in the Late Triassic ; of this diversification, however, only
29796-801: The order Plesiosauria itself. In the second half of the nineteenth century, important finds were made outside of England. While this included some German discoveries, it mainly involved plesiosaurs found in the sediments of the American Cretaceous Western Interior Seaway , the Niobrara Chalk . One fossil in particular marked the start of the Bone Wars between the rival paleontologists Edward Drinker Cope and Othniel Charles Marsh . In 1867, physician Theophilus Turner near Fort Wallace in Kansas uncovered
29987-461: The pleurapophyses decreases until they disappear between the eighth and thirteenth caudal. The height of the neural spines also decreases gradually down the tail. A row of long chevrons is present under a short portion of the tail, though not immediately behind the hips. The pectoral girdle (shoulder girdle) has a fairly standard form shared with other tanystropheids . The clavicles (collarbones) were curved and slightly twisted rods. They lie along
30178-419: The premaxilla, which extends below and behind the nares (nostril holes). The nasals (bones at the top edge of the snout) are poorly known, but were likely narrow and flat. A 2020 reinvestigation revealed that the front part of the nasals and the inner spur of the premaxillae are too short to keep the nares divided. This leaves a single central narial opening for the nostrils, opening upwards. An undivided naris
30369-444: The pterygoid). The remainder of the braincase is fully fused together into a strongly ossified composite bone, and its constituents must be estimated by comparison to other reptiles. The exoccipitals , which mostly encompass the foramen magnum (spinal cord hole), are perforated with nerve foramina. Each exoccipital merges outwards into the opisthotic , which sends out a straight, elongated paroccipital process (thick outer branch) to
30560-416: The quadrate at the rear lower corner of the skull. Nevertheless, a quadratojugal was likely present in the species, since it occurs in T. hydroides and nearly every other early archosauromorph. The paired frontals (skull roof bones above the orbits) have been described as "axe-shaped flanges", projecting broad curved plates above each orbit. Together, the frontals are narrowest at the front, terminating at
30751-411: The reptile family tree near the ancestry of archosaurs , a diverse group of reptiles with lightweight skulls and serrated teeth set in deep sockets. Dinosaurs are among the most famous subset of archosaurs, as are modern crocodilians and their prehistoric ancestors. Several newly discovered "prolacertiforms", including Tanystropheus -, Protorosaurus -, and Prolacerta -like species, were described in
30942-817: The same time, approximately 160 million years ago, the Cryptoclididae were present, shorter species with a long neck and a small head. The Leptocleididae radiated during the Early Cretaceous . These were rather small forms that, despite their short necks, might have been more closely related to the Plesiosauridae than to the Pliosauridae. Later in the Early Cretaceous, the Elasmosauridae appeared; these were among
31133-502: The seventeenth century added plesiosaur remains to their collections, such as John Woodward ; these were only much later understood to be of a plesiosaurian nature and are today partly preserved in the Sedgwick Museum . In 1719, William Stukeley described a partial skeleton of a plesiosaur, which had been brought to his attention by the great-grandfather of Charles Darwin , Robert Darwin of Elston . The stone plate came from
31324-421: The side, so they are taller than wide. Many specimens have a longitudinal lamina (ridge) on the side of each cervical. Ventral keels return to vertebrae in the rear half of the neck. All cervicals, except potentially the atlas, connected to holocephalous (single-headed) cervical ribs via facets at their front lower corner. Each cervical rib bears a short stalk connecting to two spurs running under and parallel to
31515-436: The simple fang-like premaxillary teeth, most or all of the maxillary teeth have a distinctive tricuspid shape, with the crown split into three stout triangular cusps (points). The cusps are arranged in a line from front-to-back, with the central cusp larger than the other two cusps. Among Triassic reptiles, early pterosaurs such as Eudimorphodon developed an equivalent tooth shape, and tricuspid teeth can also be found in
31706-463: The skeleton was bought by the British Museum of Natural History and catalogued as specimen NHMUK OR 22656 (formerly BMNH 22656). When the lecture was published, Conybeare also named a second species: Plesiosaurus giganteus . This was a short-necked form later assigned to the Pliosauroidea . Plesiosaurs became better known to the general public through two lavishly illustrated publications by
31897-429: The skull are rather unspecialized in this species. The vomers (front components of the palate) are narrow and dotted with at least nine tiny teeth. The succeeding palatine and pterygoid bones are also supplied with rows of teeth: up to six relatively large teeth in the former and at least 12 small teeth in the latter. Teeth on the vomers, palatines, and pterygoids are the norm for early archosauromorphs and reptiles as
32088-481: The skull). One idea was that Tanystropheus and kin (particularly Macrocnemus and Prolacerta ) were ancestral to " lacertilians ", an antequated term for lizards . This hypothesis was supported up until the 1980s by German and Swiss paleontologists, including Rupert Wild, and Peyer's successor at Zürich, Emil Kuhn-Schnyder . The other idea maintained that Tanystropheus was a "protorosaur", closer to Protorosaurus and Araeoscelis and unrelated to Prolacerta . This
32279-453: The species from Monte San Giorgio. Peyer's discoveries allowed Tribelesodon longobardicus to be recognised as a non-flying reptile, more than 40 years after its original description. Its supposed elongated finger bones were recognized as neck vertebrae, which compared favorably with those previously described as Tanystropheus from Germany and Poland. Thus, Tribelesodon longobardicus was renamed to Tanystropheus longobardicus and its anatomy
32470-417: The species have been reinterpreted as displaced postorbitals. The quadrate bone , which forms the rear edge of the skull and upper half of the jaw joint, is wide and tall. It has a strong lateral crest and a low pterygoid ramus (a vertical internal plate, articulating with the pterygoid bone in the roof of the mouth). No fossils of T. longobardicus preserve a quadratojugal , a bone which normally lies along
32661-437: The stressed vowel; the so-called breathing marks ( rough and smooth breathing ), originally used to signal presence or absence of word-initial /h/; and the diaeresis , used to mark the full syllabic value of a vowel that would otherwise be read as part of a diphthong. These marks were introduced during the course of the Hellenistic period. Actual usage of the grave in handwriting saw a rapid decline in favor of uniform usage of
32852-538: The tall suture with the postorbital and the small suture with the quadratojugal. As a result, most of the posterior skull is clustered together, and the infratemporal fenestra is reduced to a small diagonal hole. The quadratojugal is a curved sliver of bone which twists back alongside the quadrate. Relative to T. longobardicus , the quadrate has a larger pterygoid ramus and a strongly hooked projection at its upper extent. The palate of T. hydroides has several unique traits. The vomers are wide and tongue-shaped, each hosting
33043-413: The tip of the snout) has a long tooth row, with six teeth. The premaxillary teeth are conical, fluted by longitudinal ridges, and have subthecodont implantation, meaning that the inner wall of each tooth socket is lower than the outer wall. The premaxilla meets the maxilla (the succeeding toothed bone) along a long, slanted contact. This shape is produced by an elongated postnarial process (rear prong) of
33234-633: The traditional Pliosauroidea to be paraphyletic in relation to Plesiosauroidea. Rhomaleosauridae was found to be outside Neoplesiosauria, but still within Plesiosauria. The early Carnian pistosaur Bobosaurus was found to be one step more advanced than Augustasaurus in relation to the Plesiosauria and therefore it represented by definition the basalmost known plesiosaur. This analysis focused on basal plesiosaurs and therefore only one derived pliosaurid and one cryptoclidian were included, while elasmosaurids were not included at all. A more detailed analysis published by both Benson and Druckenmiller in 2014
33425-405: The turn of the century, most plesiosaur research was done by a former student of Marsh, Professor Samuel Wendell Williston . In 1914, Williston published his Water reptiles of the past and present . Despite treating sea reptiles in general, it would for many years remain the most extensive general text on plesiosaurs. In 2013, a first modern textbook was being prepared by Olivier Rieppel . During
33616-465: The underdeveloped posterior process indicates that the margin of the infratemporal fenestra (lower skull hole behind the eye) was incomplete and open from below. The postorbital bone , which links the jugal to the top of the skull, was tall and roughly boomerang-shaped, though poor preservation obscures some details. The squamosal bone , which extends behind the postorbital, is also poorly known in T. longobardicus , and many supposed squamosal fossils in
33807-571: The unusual tricuspid fossil as a pterosaur , which he named Tribelesodon longobardicus . The holotype specimen of Tribelesodon longobardicus was stored in the Museo Civico di Storia Naturale di Milano (Natural History Museum of Milan ), and was destroyed by allied bombing of Milan in World War II . Excavations by University of Zürich paleontologist Bernhard Peyer in the late 1920s and 1930s revealed many more complete fossils of
33998-450: The vertebrae. The forward-projecting spurs were short and stubby, while the rear-projecting spurs were extremely narrow and elongated, up to three times longer than their respective vertebrae. This bundle of rod-like bones running along the neck afforded a large degree of rigidity. The 12th cervical and its corresponding ribs, though still longer than tall, are notably shorter (from front-to-back) than their predecessors. The 12th cervical has
34189-461: The way they hunted and swam, incorporating general modern insights about biomechanics and ecology . The many recent discoveries have tested these hypotheses and given rise to new ones. The Plesiosauria have their origins within the Sauropterygia , a group of perhaps archelosaurian reptiles that returned to the sea. An advanced sauropterygian subgroup, the carnivorous Eusauropterygia with small heads and long necks, split into two branches during
34380-574: Was a distinct dialect of Greek itself. Aside from the Macedonian question, current consensus regards Phrygian as the closest relative of Greek, since they share a number of phonological, morphological and lexical isoglosses , with some being exclusive between them. Scholars have proposed a Graeco-Phrygian subgroup out of which Greek and Phrygian originated. Among living languages, some Indo-Europeanists suggest that Greek may be most closely related to Armenian (see Graeco-Armenian ) or
34571-568: Was also used as the official language of government and religion in the Christian Nubian kingdoms , for most of their history. Greek, in its modern form, is the official language of Greece, where it is spoken by almost the entire population. It is also the official language of Cyprus (nominally alongside Turkish ) and the British Overseas Territory of Akrotiri and Dhekelia (alongside English ). Because of
34762-557: Was also used in Ancient Greek. Greek has occasionally been written in the Latin script , especially in areas under Venetian rule or by Greek Catholics . The term Frankolevantinika / Φραγκολεβαντίνικα applies when the Latin script is used to write Greek in the cultural ambit of Catholicism (because Frankos / Φράγκος is an older Greek term for West-European dating to when most of (Roman Catholic Christian) West Europe
34953-422: Was around the same size, with the largest specimens at an estimated length of 5.25 meters (17.2 feet). T. longobardicus was significantly smaller, with an absolute maximum size of two meters (6.6 feet). Despite the large size of some Tanystropheus species, the animal was lightly built. One mass estimate used crocodiles as a density guideline for a 3.6 meter (11.8 feet)-long model of a Tanystropheus skeleton. For
35144-526: Was discovered, by a local fisherman, at Bridgwater Bay National Nature Reserve in Somerset, UK. The fossil, dating from 180 million years ago as indicated by the ammonites associated with it, measured 1.5 metres (4 ft 11 in) in length, and may be related to Rhomaleosaurus . It is probably the best preserved specimen of a plesiosaur yet discovered. In 2005, the remains of three plesiosaurs ( Dolichorhynchops herschelensis ) discovered in
35335-623: Was named as a new species, Tanystropheus hydroides (referencing the Hydra of Greek mythology ), while the smaller tricuspid morphotype retains the name T. longobardicus . The first Tanystropheus specimens to be described were found in the mid-19th century. They included eight large vertebrae from the Upper Muschelkalk of Germany , and a partial skeleton from the Lower Keuper of Poland . These geological units occupy part of
35526-423: Was originally classified as a dinosaur . The specimen is actually a very large plesiosaur, possibly reaching 15 m (49 ft) in length. The media published exaggerated reports claiming it was 25 metres (82 ft) long, and weighed up to 150,000 kilograms (330,000 lb), which would have made it among the largest predators of all time. In 2004, what appeared to be a completely intact juvenile plesiosaur
35717-579: Was popular among American paleontologists like Alfred Romer . Some publications from the mid-20th century argued that "protorosaurs" were "euryapsids" (reptiles with only an upper temporal fenestra ) related to sauropterygians, though later accounts admitted that Euryapsida was likely polyphyletic , with its members lacking a common ancestor. In 1975, a paper by South African paleontologist C.E. Gow argued that none of these hypotheses were entirely correct. He proposed that Prolacerta , and by extension Macrocnemus and Tanystropheus , occupied an extinct spur on
35908-513: Was revised into a long-necked, non-pterosaur reptile. Specimen PIMUZ T 2791, which was discovered in 1929, has been designated as the neotype of the species. Well-preserved T. longobardicus fossils continue to be recovered from Monte San Giorgio up to the present day. Fossils from the mountain are primarily stored at the rebuilt Museo Civico di Storia Naturale di Milano (MSNM), the Paleontological Museum of Zürich (PIMUZ), and
36099-420: Was the namesake of yet another term introduced into the convoluted space of reptile taxonomy: " Prolacertiformes ". As the century progressed, two competing hypotheses for the affinities of Tanystropheus developed from the groundwork set by Peyer. Both hypotheses were justified by patterns of skull fenestration (the shape of holes in the skull behind the eye) and cranial kinesis (the flexibility of joints within
36290-580: Was under the control of the Frankish Empire ). Frankochiotika / Φραγκοχιώτικα (meaning 'Catholic Chiot') alludes to the significant presence of Catholic missionaries based on the island of Chios . Additionally, the term Greeklish is often used when the Greek language is written in a Latin script in online communications. The Latin script is nowadays used by the Greek-speaking communities of Southern Italy . The Yevanic dialect
36481-450: Was written by Romaniote and Constantinopolitan Karaite Jews using the Hebrew Alphabet . In a tradition, that in modern time, has come to be known as Greek Aljamiado , some Greek Muslims from Crete wrote their Cretan Greek in the Arabic alphabet . The same happened among Epirote Muslims in Ioannina . This also happened among Arabic-speaking Byzantine rite Christians in the Levant ( Lebanon , Palestine, and Syria ). This usage
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