The Meuse or Maas is a major European river , rising in France and flowing through Belgium and the Netherlands before draining into the North Sea from the Rhine–Meuse–Scheldt delta . It has a total length of 925 km (575 miles).
111-612: Mosasaurus ( / ˌ m oʊ z ə ˈ s ɔːr ə s / ; "lizard of the Meuse River ") is the type genus (defining example) of the mosasaurs , an extinct group of aquatic squamate reptiles . It lived from about 82 to 66 million years ago during the Campanian and Maastrichtian stages of the Late Cretaceous . The genus was one of the first Mesozoic marine reptiles known to science—the first fossils of Mosasaurus were found as skulls in
222-417: A M. hoffmannii maxillary tooth measured an average rate of deposition of odontoblasts , the cells responsible for the formation of dentin , at 10.9 micrometers (0.00043 in) per day. This was by observing the von Ebner lines , incremental marks in dentin that form daily. It was approximated that it took the odontoblasts 511 days and dentin 233 days to develop to the extent observed in the tooth. One of
333-404: A 45-foot (14 m) long fish. Richard Ellis speculated in 2003 that this may have been the earliest discovery of the second species M. missouriensis , although competing speculations exist. In 1818, a fossil from Monmouth County, New Jersey became the first North American specimen to be correctly recognized as a Mosasaurus by scientists of the time. The type specimen of M. missouriensis
444-456: A chalk quarry near the Dutch city of Maastricht in the late 18th century, and were initially thought to be crocodiles or whales. One skull discovered around 1780 was famously nicknamed the "great animal of Maastricht". In 1808, naturalist Georges Cuvier concluded that it belonged to a giant marine lizard with similarities to monitor lizards but otherwise unlike any known living animal. This concept
555-835: A design specialized for cutting prey, highly prismatic surfaces (enamel circumference shaped by flat sides called prisms), and two opposite cutting edges. Mosasaurus teeth are large and robust except for those in M. conodon and M. lemonnieri , which instead have more slender teeth. The cutting edges of Mosasaurus differ by species. The cutting edges in M. hoffmannii and M. missouriensis are finely serrated, while in M. conodon and M. lemonnieri serrations do not exist. The cutting edges of M. beaugei are neither serrated nor smooth, but instead possess minute wrinkles known as crenulations. The number of prisms in Mosasaurus teeth can slightly vary between tooth types and general patterns differ between species— M. hoffmannii had two to three prisms on
666-470: A future formal reassessment. Street & Caldwell (2017) was derived from Street's 2016 doctoral thesis, which contained a phylogenetic study proposing the constraining of Mosasaurus into four species— M. hoffmannii , M. missouriensis , M. lemonnieri , and a proposed new species ' M. glycys ' —with M. conodon and the Pacific taxa recovered as belonging to different genera and M. beaugei view as
777-424: A high metabolic rate suggesting it was endothermic ("warm-blooded"), an adaptation in squamates only found in mosasaurs. There is considerable morphological variability across the currently-recognized species in Mosasaurus —from the robustly-built M. hoffmannii to the slender and serpentine M. lemonnieri —but an unclear diagnosis (description of distinguishing features) of the type species M. hoffmannii led to
888-423: A historically problematic classification. As a result, more than fifty species have been attributed to the genus in the past. A redescription of the type specimen in 2017 helped resolve the taxonomy issue and confirmed at least five species to be within the genus. Another five species still nominally classified within Mosasaurus are planned to be reassessed. Fossil evidence suggests Mosasaurus inhabited much of
999-517: A junior synonym of M. hoffmannii . As the type genus of the family Mosasauridae and the subfamily Mosasaurinae, Mosasaurus is a member of the order Squamata (which comprises lizards and snakes ). Relationships between mosasaurs and living squamates remain controversial as scientists still fiercely debate on whether the closest living relatives of mosasaurs are monitor lizards or snakes. Mosasaurus , along with mosasaur genera Eremiasaurus , Plotosaurus , and Moanasaurus traditionally form
1110-582: A juvenile form of M. hoffmannii based on the argument that significant differences could be explained by age-based variation. However, the need for more research to confirm any hypotheses of synonymy was expressed. The fifth species M. beaugei was described by Camille Arambourg in 1952 from isolated teeth originating from phosphate deposits in the Oulad Abdoun Basin and the Ganntour Basin in Morocco,
1221-578: A maximum length of 17.1 meters (56 ft). Using a smaller partial jaw ( NHMM 009002) measuring 90 centimeters (35 in) and "reliably estimated at" 160 centimeters (63 in) when complete, Lingham-Soliar (1995) estimated a larger maximum length of 17.6 meters (58 ft) via the same ratio. No explicit justification for the 1:10 ratio was provided in Russell (1967), and it has been considered to be probably overestimated by Cleary et al. (2018). In 2014, Federico Fanti and colleagues alternatively argued that
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#17328551775831332-571: A paddle. Schlegel's hypothesis was largely ignored by contemporary scientists but became widely accepted by the 1870s when Othniel Charles Marsh and Cope uncovered more complete mosasaur remains in North America. One of the earliest depictions of Mosasaurus in paleoart is a life-size concrete sculpture created by Benjamin Waterhouse Hawkins between 1852 and 1854 as part of the collection of sculptures of prehistoric animals on display at
1443-676: A proper diagnosis during its initial descriptions, which led to ambiguity in how the genus is defined. This led Mosasaurus to become a wastebasket taxon containing as many as fifty different species. A 2017 study by Hallie Street and Michael Caldwell performed the first proper diagnosis and description of the M. hoffmannii holotype, which allowed a major taxonomic cleanup confirming five species as likely valid— M. hoffmannii , M. missouriensis , M. conodon , M. lemonnieri , and M. beaugei . The study also held four additional species from Pacific deposits— M. mokoroa , M. hobetsuensis , M. flemingi , and M. prismaticus —to be possibly valid, pending
1554-450: A sister relationship with another group which included Globidens and Prognathodon , and M. maximus as a sister species to Plotosaurus . The latter rendered Mosasaurus paraphyletic (an unnatural grouping), but Bell (1997) nevertheless recognized Plotosaurus as a distinct genus. Bell's study served as a precedent for later studies that mostly left the systematics of Mosasaurus unchanged, although some later studies have recovered
1665-425: A skull approaching 1 meter (3.3 ft) in length. Based on personal observations of various unpublished fossils from Morocco, Nathalie Bardet et al. (2015) estimated that M. beaugei grew to a total length of 8–10 meters (26–33 ft), their skulls typically measuring around 1 meter (3.3 ft) in length. With a referred skull measuring 97.7 centimeters (38.5 in) in length, M. conodon has been regarded as
1776-450: A small to medium-sized representative of the genus. The skull of Mosasaurus is conical and tapers off to a short snout which extends a little beyond the frontmost teeth. In M. hoffmannii , this snout is blunt, while in M. lemonnieri it is pointed. Above the gum line in both jaws, a single row of small pits known as foramina are lined parallel to the jawline; they are used to hold the terminal branches of jaw nerves. The foramina along
1887-600: A species of Mosasaurus . Although Cope did not provide the etymology for the specific epithet conodon , it is suggested that it could be a portmanteau meaning "conical tooth", derived from the Ancient Greek κῶνος ( kônos , "cone") and ὀδών ( odṓn , "tooth"), probably in reference to conical surface teeth smooth of the species. The fourth species M. lemonnieri was first detected by Camper Jr. based on fossils from his father's collections, which he discussed with Cuvier during their 1799 correspondence. Although Cuvier rejected
1998-639: A tribe within the Mosasaurinae variously called Mosasaurini or Plotosaurini. One of the earliest relevant attempts at an evolutionary study of Mosasaurus was done by Russell in 1967. He proposed that Mosasaurus evolved from a Clidastes -like mosasaur, and diverged into two lineages, one giving rise to M. conodon and another siring a chronospecies sequence which contained in order of succession M. ivoensis , M. missouriensis , and M. maximus-hoffmanni . However, Russell used an early method of phylogenetics and did not use cladistics. In 1997, Bell published
2109-633: Is mined from chalk deposits both above ground and underground . Chalk mining boomed during the Industrial Revolution , due to the need for chalk products such as quicklime and bricks . Most people first encounter chalk in school where it refers to blackboard chalk , which was originally made of mineral chalk, since it readily crumbles and leaves particles that stick loosely to rough surfaces, allowing it to make writing that can be readily erased. Blackboard chalk manufacturers now may use mineral chalk, other mineral sources of calcium carbonate, or
2220-610: Is a fine-textured, earthy type of limestone distinguished by its light colour, softness, and high porosity. It is composed mostly of tiny fragments of the calcite shells or skeletons of plankton , such as foraminifera or coccolithophores . These fragments mostly take the form of calcite plates ranging from 0.5 to 4 microns in size, though about 10% to 25% of a typical chalk is composed of fragments that are 10 to 100 microns in size. The larger fragments include intact plankton skeletons and skeletal fragments of larger organisms, such as molluscs , echinoderms , or bryozoans . Chalk
2331-404: Is a soft, white, porous , sedimentary carbonate rock . It is a form of limestone composed of the mineral calcite and originally formed deep under the sea by the compression of microscopic plankton that had settled to the sea floor . Chalk is common throughout Western Europe , where deposits underlie parts of France, and steep cliffs are often seen where they meet the sea in places such as
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#17328551775832442-508: Is also the only form of limestone that commonly shows signs of compaction. Flint (a type of chert ) is very common as bands parallel to the bedding or as nodules in seams , or linings to fractures , embedded in chalk. It is probably derived from sponge spicules or other siliceous organisms as water is expelled upwards during compaction. Flint is often deposited around larger fossils such as Echinoidea which may be silicified (i.e. replaced molecule by molecule by flint). Chalk
2553-428: Is applied to the hands and feet to remove perspiration and reduce slipping. Chalk may also be used as a house construction material instead of brick or wattle and daub : quarried chalk was cut into blocks and used as ashlar , or loose chalk was rammed into blocks and laid in mortar. There are still houses standing which have been constructed using chalk as the main building material. Most are pre- Victorian though
2664-433: Is around 1.5 times longer than the femur . The femur itself is about twice as long as it is wide and ends at the distal side in a pair of distinct articular facets (of which one connects to the ilium and the other to the paddle bones) that meet at an angle of approximately 120°. Five sets of metacarpals and phalanges (finger bones) were encased in and supported the paddles, with the fifth set being shorter and offset from
2775-518: Is commonly used as a drying agent to obtain better grip by gymnasts and rock climbers. Glazing putty mainly contains chalk as a filler in linseed oil . Chalk and other forms of limestone may be used for their properties as a base . Chalk is a source of quicklime by thermal decomposition , or slaked lime following quenching of quicklime with water. In agriculture , chalk is used for raising pH in soils with high acidity . Small doses of chalk can also be used as an antacid . Additionally,
2886-485: Is estimated to measure up to 12 meters (39 ft) in maximum length, making it one of the largest mosasaurs. The skull of Mosasaurus had robust jaws and strong muscles capable of powerful bites using dozens of large teeth adapted for cutting prey . Its four limbs were shaped into paddles to steer the animal underwater. Its tail was long and ended in a downward bend and a paddle-like fluke. Mosasaurus possessed excellent vision to compensate for its poor sense of smell, and
2997-571: Is now usually made of talc (magnesium silicate). Chalk beds form important petroleum reservoirs in the North Sea and along the Gulf Coast of North America. In southeast England, deneholes are a notable example of ancient chalk pits. Such bell pits may also mark the sites of ancient flint mines, where the prime object was to remove flint nodules for stone tool manufacture. The surface remains at Cissbury are one such example, but perhaps
3108-489: Is one of the largest marine reptiles known, though knowledge of its skeleton remains incomplete as it is mainly known from skulls. Russell (1967) wrote that the length of the jaw equalled one tenth of the body length in the species. Based on this ratio, Grigoriev (2014) used the largest lower jaw attributed to M. hoffmannii (CCMGE 10/2469, also known as the Penza specimen; measuring 171 centimeters (67 in) in length) to estimate
3219-586: Is so common in Cretaceous marine beds that the Cretaceous Period was named for these deposits. The name Cretaceous was derived from Latin creta , meaning chalk . Some deposits of chalk were formed after the Cretaceous. The Chalk Group is a European stratigraphic unit deposited during the late Cretaceous Period. It forms the famous White Cliffs of Dover in Kent , England, as well as their counterparts of
3330-401: Is typically almost pure calcite, CaCO 3 , with just 2% to 4% of other minerals. These are usually quartz and clay minerals , though collophane (cryptocrystalline apatite , a phosphate mineral) is also sometimes present, as nodules or as small pellets interpreted as fecal pellets. In some chalk beds, the calcite has been converted to dolomite , CaMg(CO 3 ) 2 , and in a few cases
3441-723: The Cap Blanc Nez on the other side of the Dover Strait . The Champagne region of France is mostly underlain by chalk deposits, which contain artificial caves used for wine storage . Some of the highest chalk cliffs in the world occur at Jasmund National Park in Germany and at Møns Klint in Denmark . Chalk deposits are also found in Cretaceous beds on other continents, such as the Austin Chalk , Selma Group , and Niobrara Formations of
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3552-561: The Crystal Palace Park in London . The restoration was primarily informed by Richard Owen 's interpretation of the M. hoffmannii holotype and the anatomy of monitor lizards, so Hawkins depicted the animal as essentially a water-going monitor lizard. It was given a boxy head, nostrils at the side of the skull, large volumes of soft tissue around the eyes, lips reminiscent of monitor lizards, scales consistent with those in large monitors like
3663-525: The Dover cliffs on the Kent coast of the English Channel . Chalk is mined for use in industry, such as for quicklime , bricks and builder's putty , and in agriculture , for raising pH in soils with high acidity . It is also used for " blackboard chalk " for writing and drawing on various types of surfaces, although these can also be manufactured from other carbonate-based minerals, or gypsum . Chalk
3774-520: The Komodo dragon , and a flipper. The model was deliberately sculpted incomplete, which Mark Witton believed was likely to save time and money. Many elements of the sculpture can be considered inaccurate, even for the time. It did not take into account Golduss' 1845 study of M. missouriensis which instead called for a narrower skull, nostrils at the top of the skull, and amphibious terrestrial limbs (the latter being incorrect in modern standards). Mosasaurus
3885-764: The Langres plateau in France from where it flows northwards past Sedan (the head of navigation ) and Charleville-Mézières into Belgium. At Namur it is joined by the Sambre . Beyond Namur the Meuse winds eastwards and passes Liège before turning north. The river then forms part of the Belgian-Dutch border, except that at Maastricht the border lies further to the west. In the Netherlands it continues northwards through Venlo closely along
3996-500: The M. missouriensis holotype, which indicated an elastic vertebral column that Goldfuss in 1845 saw as evidence of an ability to walk and interpretations of some phalanges as claws. In 1854, Hermann Schlegel proved how Mosasaurus actually had fully aquatic flippers. He clarified that earlier interpretations of claws were erroneous and demonstrated how the phalanges show no indication of muscle or tendon attachment, which would make walking impossible. They are also broad, flat, and form
4107-667: The Oude Maasje . In the late 19th century and early 20th century the connection between the Maas and Rhine was closed off and the Maas was given a new, artificial mouth – the Bergse Maas . The resulting separation of the rivers Rhine and Maas reduced the risk of flooding and was considered to be the greatest achievement in Dutch hydraulic engineering before the completion of the Zuiderzee Works and Delta Works . The former main branch was, after
4218-557: The Pembina Gorge State Recreation Area in North Dakota was found to have an unusual count of sixteen pterygoid teeth, far greater than in known species. The dentition was thecodont (tooth roots deeply cemented within the jaw bone). Teeth were constantly shed through a process where the replacement tooth developed within the root of the original tooth and then pushed it out of the jaw. Chemical studies conducted on
4329-587: The battle , which saw the men from Liège defeated, John ordered the drowning in the Meuse of burghers and noblemen in Liège whose loyalties he suspected. The border remained relatively stable until the annexation of the Three Bishoprics Metz, Toul and Verdun by King Henry II in 1552 and the occupation of the Duchy of Lorraine by the forces of King Louis XIII in 1633. Its lower Belgian ( Walloon ) portion, part of
4440-450: The sillon industriel , was the first fully industrialized area in continental Europe. The Afgedamde Maas was created in the late Middle Ages, when a major flood made a connection between the Maas and the Merwede at the town of Woudrichem . From that moment on, the current Afgedamde Maas was the main branch of the lower Meuse. The former main branch eventually silted up and is today called
4551-743: The Atlantic Ocean and the adjacent seaways. Mosasaurus fossils have been found in North and South America, Europe, Africa, Western Asia, and Antarctica. This distribution encompassed a wide range of oceanic climates including tropical, subtropical, temperate, and subpolar. Mosasaurus was a common large predator in these oceans and was positioned at the top of the food chain . Paleontologists believe its diet would have included virtually any animal; it likely preyed on bony fish, sharks, cephalopods , birds, and other marine reptiles including sea turtles and other mosasaurs. It likely preferred to hunt in open water near
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4662-632: The Meuse basin was one of the many regions in Europe to experience catastrophic flooding during the 2021 European floods . The name Meuse is derived from the French name of the river, derived from its Latin name, Mosa , which ultimately derives from the Celtic or Proto-Celtic name * Mosā . This probably derives from the same root as English " maze ", referring to the river's twists and turns. The Dutch name Maas descends from Middle Dutch Mase , which comes from
4773-597: The Meuse but pumps running water into the Meuse). Most of the basin area (approximately 36,000 km ) is in Wallonia (12,000 km ), followed by France (9,000 km ), the Netherlands (8,000 km ), Germany (2,000 km ), Flanders (2,000 km ) and Luxembourg (a few km ). An International Commission on the Meuse has the responsibility of the implementation of the treaty. The costs of this Commission are met by all these countries, in proportion of their own territory in
4884-440: The Meuse has been relatively stable over the last few thousand years. One recent study estimates that average flow has increased by about 10% since 2000 BC. The hydrological distribution of the Meuse changed during the later Middle Ages, when a major flood forced it to shift its main course northwards towards the river Merwede . From then on several stretches of the original Merwede were renamed "Maas" (i.e. Meuse) and served as
4995-418: The Meuse is bypassed by the 36 km (22.4 mi) Juliana Canal . South of Namur, further upstream, the river can only carry more modest vessels, although a barge as long as 100 m (328 ft). can still reach the French border town of Givet. From Givet, the river is canalized over a distance of 272 kilometres (169 mi). The canalized Meuse used to be called the "Canal de l'Est — Branche Nord" but
5106-509: The Meuse", in reference to the river where the holotype specimen was discovered nearby. In 1829, Gideon Mantell added the specific epithet hoffmannii , in honor to Hoffmann. Later, the second skull is designated as the new species' holotype (defining example). In 1804, the Lewis and Clark Expedition discovered a now-lost fossil skeleton alongside the Missouri River , which was identified as
5217-537: The Mosasaurinae, with proposed new taxa and renamings in single quotations. Dallasaurus turneri Clidastes liodontus Clidastes moorevillensis Clidastes propython Prognathodon overtoni Prognathodon rapax Prognathodon saturator Prognathodon currii Prognathodon solvayi Prognathodon waiparaensis Prognathodon kianda Eremiasaurus heterodontus Plesiotylosaurus crassidens Mosasaurus conodon Mosasaurus missouriensis Meuse (river) From 1301,
5328-411: The Netherlands, which despite the famous discovery of the M. hoffmannii holotype attracted little attention from mosasaur paleontologists. Second, the species was overshadowed by the more famous and history-rich type species . M. lemonnieri is a controversial taxon, and there is debate on whether it is a distinct species or not. In 1967, Dale Russell argued that M. lemonnieri and M. conodon are
5439-698: The North American interior. Chalk is also found in western Egypt (Khoman Formation) and western Australia ( Miria Formation ). Chalk of Oligocene to Neogene age has been found in drill cores of rock under the Pacific Ocean at Stewart Arch in the Solomon Islands . There are layers of chalk, containing Globorotalia , in the Nicosia Formation of Cyprus , which formed during the Pliocene . Chalk
5550-562: The Rhine distributaries. The resulting separation of the rivers Rhine and Meuse is considered to be the greatest achievement in Dutch hydraulic engineering before the completion of the Zuiderzee Works and Delta Works . In 1970 the Haringvlietdam has been finished. Since then the reunited Rhine and Meuse waters have reached the North Sea either at this site or, during times of lower discharges of
5661-412: The Rhine, at Hook of Holland . A 2008 study notes that the difference between summer and winter flow volumes has increased significantly in the last 100–200 years. It points out that the frequency of serious floods ( i.e. flows > 1000% of normal) has increased markedly. They predict that winter flooding of the Meuse may become a recurring problem in the coming decades. The Meuse flows through
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#17328551775835772-627: The Waal, the main stem of the Rhine at Woudrichem , and then flows under the name of Boven Merwede to Hardinxveld-Giessendam , where it splits into Nieuwe Merwede and Beneden Merwede . Near Lage Zwaluwe , the Nieuwe Merwede joins the Amer, forming the Hollands Diep , which splits into Grevelingen and Haringvliet , before finally flowing into the North Sea. The Meuse is crossed by railway bridges between
5883-579: The basin of the Meuse: Netherlands 30%, Wallonia 30%, France 15%, Germany 14.5%, Flanders 5%, Brussels 4.5%, Kingdom of Belgium 0.5%, and Luxembourg 0.5%. The map of the basin area of Meuse was joined to the text of the treaty. As for culture, as a major communication route the River Meuse is the origin of Mosan art , principally (Wallonia and France). The first landscape painted in the Renaissance
5994-404: The bones is virtually identical with in modern whales, which indicates Mosasaurus possessed a high range of aquatic adaptation and neutral buoyancy as seen in cetaceans. The tail structure of Mosasaurus is similar to relatives like Prognathodon , in which soft tissue evidence for a two-lobed tail is known. The tail vertebrae gradually shorten around the center of the tail and lengthen behind
6105-644: The border to Germany, then turns towards the west, where it runs parallel to the Waal and forms part of the extensive Rhine–Meuse–Scheldt delta , together with the Scheldt to its south and the Rhine to the north. The river has been divided near Heusden into the Afgedamde Maas on the right and the Bergse Maas on the left. The Bergse Maas continues under the name of Amer , which is part of De Biesbosch . The Afgedamde Maas joins
6216-506: The case with the largest specimens of M. lemonnieri , although more typical skulls of the species have a near-perfectly straight jawline. The premaxillary bar, the long portion of the premaxillary bone extending behind the premaxillary teeth, is narrow and constricts near the middle in M. hoffmannii and M. lemonnieri like in typical mosasaurs. In M. missouriensis , the bar is robust and does not constrict. The external nares ( nostril openings ) are moderately sized and measure around 21–24% of
6327-399: The center, suggesting rigidness around the tail center and excellent flexibility behind it. Like most advanced mosasaurs, the tail bends slightly downwards as it approached the center, but this bend is offset from the dorsal plane at a small degree. Mosasaurus also has large haemal arches located at the bottom of each caudal vertebra which bend near the middle of the tail, which contrasts with
6438-548: The chalk came mostly form low-magnesium calcite skeletons, so the sediments were already in the form of highly stable low-magnesium calcite when deposited. This is in contrast with most other limestones, which formed from high-magnesium calcite or aragonite that rapidly converted to the more stable low-magnesium calcite after deposition, resulting in the early cementation of such limestones. In chalk, absence of this calcium carbonate conversion process prevented early cementation, which partially accounts for chalk's high porosity. Chalk
6549-684: The dam at its southern inlet was completed in 1904, renamed Afgedamde Maas and no longer receives water from the Maas. The Meuse and its crossings were a key objective of the Battle of France , the Battle of Sedan and also for the last major German WWII counter-offensive on the Western Front , the Battle of the Bulge in December 1944 and January 1945. The Meuse is represented in the documentary The River People released in 2012 by Xavier Istasse. In July 2021,
6660-485: The dolomitized chalk has been dedolomitized back to calcite. Chalk is highly porous, with typical values of porosity ranging from 35 to 47 per cent. While it is similar in appearance to both gypsum and diatomite , chalk is identifiable by its hardness, fossil content, and its reaction to acid (it produces effervescence on contact). In Western Europe, chalk was formed in the Late Cretaceous Epoch and
6771-437: The early Palaeocene Epoch (between 100 and 61 million years ago). It was deposited on extensive continental shelves at depths between 100 and 600 metres (330 and 1,970 ft), during a time of nonseasonal (likely arid) climate that reduced the amount of erosion from nearby exposed rock. The lack of nearby erosion explains the high purity of chalk. The coccolithophores, foraminifera, and other microscopic organisms from which
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#17328551775836882-420: The first cladistical study of North American mosasaurs. Incorporating the species M. missouriensis , M. conodon , M. maximus , and an indeterminate specimen ( UNSM 77040), some of his findings agreed with Russell (1967), such as Mosasaurus descending from an ancestral group containing Clidastes and M. conodon being the most basal of the genus. Contrary to Russell (1967), Bell also recovered Mosasaurus in
6993-1058: The following departments of France , provinces of Belgium , provinces of the Netherlands and towns: Main cities and tributaries will be in bold . France Grand Est Region Haute-Marne Department [REDACTED] Le Châtelet-sur-Meuse [REDACTED] Premier pont de la Meuse [REDACTED] Pont de Malroy [REDACTED] Pont du Pâtis des Vannees [REDACTED] Ruisseau de Pré Chatenay [REDACTED] Pont de Meuse (D429 Val-de-Meuse - Dombrot-le-Sec ) [REDACTED] Ruisseau d'Avrecourt [REDACTED] Railway bridge Culmont-Chalindrey - Toul line [REDACTED] Ru d'Ouette [REDACTED] Ru des Fossés [REDACTED] Ruisseau de Bocheret [REDACTED] Provenchères-sur-Meuse [REDACTED] Pont de Val-de-Meuse (D189) [REDACTED] Ruisseau des Aimeguenons [REDACTED] Pont de l'A31 ( A31 Nancy - Dijon ) [REDACTED] Ruisseau de Joncourt [REDACTED] Pont de D132 [REDACTED] Ruisseau de l'Étange Chalk Chalk
7104-575: The following stations (on the left and right banks respectively): There are also numerous road bridges and around 32 ferry crossings. The Meuse is navigable over a substantial part of its total length: In the Netherlands and Belgium, the river is part of the major inland navigation infrastructure, connecting the Rotterdam-Amsterdam-Antwerp port areas to the industrial areas upstream: 's-Hertogenbosch, Venlo, Maastricht, Liège, Namur. Between Maastricht and Maasbracht , an unnavigable section of
7215-562: The fossil into cultural fame, but historians agree that the narrative was exaggerated. After its seizure, the second skull was sent to the National Museum of Natural History, France in 1795 and later cataloged as MNHN AC 9648. By 1800, Camper's son Adriaan Gilles Camper concluded that the fossil, which by then was nicknamed the "great animal of Maastricht", belonged to a marine reptile sharing affinities to monitor lizards , but otherwise unlike any modern animal. Georges Cuvier confirmed
7326-645: The genus's paraphyletic status. Third, there was still a lack of comparative studies of the skeletal anatomy of large mosasaurines at the time. These problems were addressed in Street's 2016 thesis in an updated phylogenetic analysis. Conrad uniquely used only M. hoffmannii and M. lemonnieri in his 2008 phylogenetic analysis, which recovered M. hoffmannii as basal to a multitude of descendant clades containing (in order of most to least basal) Globidens , M. lemonnieri , Goronyosaurus , and Plotosaurus . This result indicated that M. hoffmannii and M. lemonnieri are not in
7437-472: The holotype tooth being cataloged as MNHN PMC 7. The species is named in honor of Alfred Beaugé, director at the time of the OCP Group , who invited Arambourg to participate in the research project and helped him to provide local fossils. Scientists during the early and mid-1800s initially imagined Mosasaurus as an amphibious marine reptile with webbed feet and limbs for walking . This was based on fossils like
7548-458: The idea of another Mosasaurus species, Camper Jr. nevertheless published his findings in 1812 without establishing a scientific name. This species was re-introduced to science and formally described in 1889 by Louis Dollo based on a skull recovered by Alfred Lemonnier from a phosphate quarry in Belgium, a holotype since cataloged as IRSNB R28. Dollo names the species in his honor. Further mining of
7659-859: The jaws ( homodont ) except for the smaller pterygoid teeth. The number of teeth in the maxillae, pterygoids, and dentaries vary between species and sometimes even individuals— M. hoffmannii had fourteen to sixteen maxillary teeth, fourteen to fifteen dentary teeth, and eight pterygoid teeth; M. missouriensis had fourteen to fifteen maxillary teeth, fourteen to fifteen dentary teeth, and eight to nine pterygoid teeth; M. conodon had fourteen to fifteen maxillary teeth, sixteen to seventeen dentary teeth, and eight pterygoid teeth; M. lemonnieri had fifteen maxillary teeth, fourteen to seventeen dentary teeth, and eleven to twelve pterygoid teeth; and M. beaugei had twelve to thirteen maxillary teeth, fourteen to sixteen dentary teeth, and six or more pterygoid teeth. One indeterminate specimen of Mosasaurus similar to M. conodon from
7770-431: The labial side (the side facing outwards) and no prisms on the lingual side (the side facing the tongue), M. missouriensis had four to six labial prisms and eight lingual prisms, M. lemonnieri had eight to ten labial prisms, and M. beaugei had three to five labial prisms and eight to nine lingual prisms. Like all mosasaurs, Mosasaurus had four types of teeth, classified based on the jaw bones they were located on. On
7881-445: The latter. Spaces within the braincase for the occipital lobe and cerebral hemisphere are narrow and shallow, suggesting such brain parts were relatively small. The parietal foramen in Mosasaurus , which is associated with the parietal eye , is the smallest among mosasaurids . The quadrate bone, which connected the lower jaw to the rest of the skull and formed the jaw joint, is tall and somewhat rectangular in shape, differing from
7992-409: The left (Topology A) is modified from a maximum clade credibility tree inferred by a Bayesian analysis in the most recent major phylogenetic analysis of the Mosasaurinae subfamily by Madzia & Cau (2017), which was self-described as a refinement of a larger study by Simões et al. (2017). The cladogram on the right (Topology B) is modified from Street's 2016 doctoral thesis proposing a revision to
8103-528: The mineral gypsum ( calcium sulfate ). While gypsum-based blackboard chalk is the lowest cost to produce, and thus widely used in the developing world , use of carbonate-based chalk produces larger particles and thus less dust, and it is marketed as "dustless chalk". Coloured chalks, pastel chalks, and sidewalk chalk (shaped into larger sticks and often coloured), used to draw on sidewalks , streets, and driveways , are primarily made of gypsum rather than calcium carbonate chalk. Magnesium carbonate chalk
8214-881: The most complete Mosasaurus skeletons in terms of vertebral representation ( Mosasaurus sp.; SDSM 452) has seven cervical (neck) vertebrae , thirty-eight dorsal vertebrae (which includes thoracic and lumbar vertebrae ) in the back, and eight pygal vertebrae (front tail vertebrae lacking haemal arches ) followed by sixty-eight caudal vertebrae in the tail. All species of Mosasaurus have seven cervical vertebrae, but other vertebral counts vary among them. Various partial skeletons of M. conodon , M. hoffmannii , and M. missouriensis suggest M. conodon likely had up to thirty-six dorsal vertebrae and nine pygal vertebrae; M. hoffmannii had likely up to thirty-two dorsal vertebrae and ten pygal vertebrae; and M. missouriensis around thirty-three dorsal vertebrae, eleven pygal vertebrae, and at least seventy-nine caudal vertebrae. M. lemmonieri had
8325-557: The most famous is the extensive complex at Grimes Graves in Norfolk . Chalk was traditionally used in recreation. In field sports, such as tennis played on grass, powdered chalk was used to mark the boundary lines of the playing field or court. If a ball hits the line, a cloud of chalk or pigment dust will be visible. In recent years, powdered chalk has been replaced with titanium dioxide . In gymnastics, rock-climbing, weightlifting and tug of war , chalk — now usually magnesium carbonate —
8436-485: The most vertebrae in the genus, with up to around forty dorsal vertebrae, twenty-two pygal vertebrae, and ninety caudal vertebrae. Compared to other mosasaurs, the rib cage of Mosasaurus is unusually deep and forms an almost perfect semicircle, giving it a barrel-shaped chest. Rather than being fused together, extensive cartilage likely connected the ribs with the sternum , which would have facilitated breathing movements and compression when in deeper waters. The texture of
8547-476: The observations of Camper Jr. in a more in-depth study which was published in 1808. The skull became part of Cuvier's first speculations about the conception of extinction , which later led to his theory of catastrophism , a precursor to the theory of evolution . At the time, it was not believed that a species could go extinct, and fossils of animals were often interpreted as some form of an extant species. Cuvier's idea that there existed an animal unlike any today
8658-500: The possession of prince Maximilian of Weid-Neuwied between 1832 and 1834. The fossil skull, now cataloged as RFWUIP 1327, was delivered to Georg August Goldfuss in Bonn for research, who published a study in 1845. The same year, Christian Erich Hermann von Meyer suspected that the skull and Harlan's snout were part of the same individual. This could not be confirmed at the time because the fossil snout became lost. The von Meyer's suggestion
8769-596: The presumed but unattested Old Dutch form * Masa , from Proto-Germanic * Masō . Modern Dutch and German Maas and Limburgish Maos preserve this Germanic form. Despite the similarity, the Germanic name is not derived from the Celtic name, judging from the change from earlier o into a , which is characteristic of the Germanic languages. The Meuse rises in Pouilly-en-Bassigny, commune of Le Châtelet-sur-Meuse on
8880-580: The primary outflow of that river. Those branches are currently known as the Nieuwe Maas and Oude Maas. However during another series of severe floods the Meuse found an additional path towards the sea, resulting in the creation of the Biesbosch wetlands and Hollands Diep estuaries. Thereafter the Meuse split near Heusden into two main distributaries, one flowing north to join the Merwede and one flowing direct to
8991-575: The prominent biologist Petrus Camper , and the skull gained international attention after Camper published a study identifying it as a whale. This caught the attention of French revolutionaries , who looted the fossil following the capture of Maastricht during the French Revolutionary Wars in 1794. In a 1799 narrative of this event by Barthélemy Faujas de Saint-Fond , the skull was allegedly retrieved by twelve grenadiers in exchange for an offer of 600 bottles of wine. This story helped elevate
9102-411: The pterygoid bones, palatine bone , and nearby processes of other bones, is tightly packed to provide greater cranial stability. The neurocranium housed a brain which was narrow and relatively small compared to other mosasaurs. For example, the braincase of the mosasaur Plioplatecarpus marshi provided for a brain around twice the size of that in M. hoffmannii despite being only half the length of
9213-487: The quarry in subsequent years uncovered many additional well-preserved fossils, including multiple partial skeletons which collectively represented nearly the entire skeleton of the species. They were described by Dollo in later papers. Despite being one of the best anatomically represented species, M. lemonnieri was largely ignored in scientific literature. Theagarten Lingham-Soliar suggested two reasons for this neglect. First, M. lemonnieri fossils are endemic to Belgium and
9324-436: The reduction of haemal arches in other marine reptiles such as ichthyosaurs . These and other features support a large and powerful paddle-like fluke in Mosasaurus . The forelimbs of Mosasaurus are wide and robust. The scapula and humerus are fan-shaped and wider than tall. The radius and ulna are short, but the former is taller and larger than the latter. The ilium is rod-like and slender; in M. missouriensis , it
9435-504: The rest. The overall structure of the paddle is compressed, similar to in Plotosaurus , and was well-suited for faster swimming. In the hindlimbs, the paddle is supported by four sets of digits. Interactive skeletal reconstruction of M. hoffmannii (hover over or click on each skeletal component to identify the structure) Because nomenclatural rules were not well-defined at the time, 19th century scientists did not give Mosasaurus
9546-499: The river Meuse. The first fossils of it were discovered outside Maastricht in 1780. An international agreement was signed in 2002 in Ghent , Belgium, about the management of the river amongst France, Germany, Luxembourg , the Netherlands, and Belgium. Also participating in the agreement were the Belgian regional governments of Flanders , Wallonia , and Brussels (which is not in the basin of
9657-509: The rounder quadrates found in typical mosasaurs. The quadrate also housed the hearing structures , with the eardrum residing within a round and concave depression in the outer surface called the tympanic ala. The trachea likely stretched from the esophagus to below the back end of the lower jaw's coronoid process , where it split into smaller pairs of bronchi which extended parallel to each other. The features of teeth in Mosasaurus vary across species, but unifying characteristics include
9768-410: The same ecosystems through niche partitioning . There were still conflicts among them, as an instance of Tylosaurus attacking a Mosasaurus has been documented. Several fossils document deliberate attacks on Mosasaurus individuals by members of the same species. In fighting likely took place in the form of snout grappling, as seen in modern crocodiles. The first Mosasaurus fossil known to science
9879-462: The same genus. However, the study used a method unorthodox to traditional phylogenetic studies on mosasaur species because its focus was on the relationships of entire squamate groups rather than mosasaur classification. As a result, some paleontologists caution that lower-order classification results from Conrad's 2008 study such as the specific placement of Mosasaurus may contain technical problems, making them inaccurate. The following cladogram on
9990-489: The same ratio, Gayford et al. (2024) calculated the total length for the Penza specimen to be 12 meters (39 ft). Isolated bones suggest some M. hoffmannii may have exceeded the lengths of the Penza specimen. One such bone is a quadrate (NHMM 003892) which is 150% larger than the average size, which Everhart and colleagues in 2016 reported can be extrapolated to scale an individual around 18 meters (59 ft) in length. It
10101-417: The same species and designated the former as a junior synonym per the principle of priority . In a 2000 study, Lingham-Soliar refuted this based on a comprehensive study of existing M. lemonnieri specimens, which was corroborated by a study on the M. conodon skull by Takehito Ikejiri and Spencer G. Lucas in 2014. In 2004, Eric Mulder, Dirk Cornelissen, and Louis Verding suggested M. lemonnieri could be
10212-413: The sea. The branch of the Meuse leading direct to the sea eventually silted up (and now forms the Oude Maasje stream), but in 1904 the canalised Bergse Maas was dug to take over the functions of the silted-up branch. At the same time the branch leading to the Merwede was dammed at Heusden (and has since been known as the Afgedamde Maas ) so that little water from the Meuse entered the old Maas courses or
10323-449: The sister group to Mosasaurus and Plotosaurus to instead be Eremiasaurus or Plesiotylosaurus depending on the method of data interpretation used, with at least one study also recovering M. missouriensis to be the most basal species of the genus instead of M. conodon . In 2014, Konishi and colleagues expressed a number of concerns with the reliance on Bell's study. First, the genus was severely underrepresented by incorporating only
10434-442: The skull constituted approximately one-eleventh of the whole body. Polcyn et al. (2014) estimated that M. missouriensis may have measured up to 8–9 meters (26–30 ft) in length. Street (2016) noted that large M. missouriensis individuals typically had skulls exceeding lengths of 1 meter (3.3 ft). A particular near-complete skeleton of M. missouriensis is reportedly measured at 6.5 meters (21 ft) in total length with
10545-422: The skull's length in M. hoffmannii . They are placed further toward the back of the skull than in nearly all other mosasaurs (exceeded only by Goronyosaurus ), and begin above the fourth or fifth maxillary teeth. As a result, the rear portions of the maxilla (the main tooth-bearing bone of the upper jaw) lack the dorsal concavity that would fit the nostrils in typical mosasaurs. The palate , which consists of
10656-436: The small particles of chalk make it a substance ideal for cleaning and polishing. For example, toothpaste commonly contains small amounts of chalk, which serves as a mild abrasive . Polishing chalk is chalk prepared with a carefully controlled grain size, for very fine polishing of metals. French chalk (also known as tailor's chalk) is traditionally a hard chalk used to make temporary markings on cloth, mainly by tailors . It
10767-453: The snout form a pattern similar to the foramina in Clidastes skulls. The upper jaws in most species are robustly built, broad, and deep except in M. conodon , where they are slender. The disparity is also reflected in the dentary , the lower jawbone, although all species share a long and straight dentary. In M. hoffmannii , the top margin of the dentary is slightly curved upwards; this is also
10878-541: The surface. From an ecological standpoint, Mosasaurus probably had a profound impact on the structuring of marine ecosystems; its arrival in some locations such as the Western Interior Seaway in North America coincides with a complete turnover of faunal assemblages and diversity. Mosasaurus faced competition with other large predatory mosasaurs such as Prognathodon and Tylosaurus —which were known to feed on similar prey—though they were able to coexist in
10989-417: The three North American species M. hoffmannii/M. maximus , M. missouriensis , and M. conodon ; by doing so, others like M. lemonnieri , which is one of the most completely known species in the genus, were neglected, which affected phylogenetic results. Second, the studies relied on an unclean and shaky taxonomy of the Mosasaurus genus due to the lack of a clear holotype diagnosis, which may have been behind
11100-436: The total length of M. hoffmannii was more likely closer to seven times the length of the skull, which was based on a near-complete skeleton of the related species Prognathodon overtoni . The study estimated that an M. hoffmannii individual with a skull measuring more than 145 cm (57 in) would have been up to or more than 11 meters (36 ft) in length and weighed 10 metric tons (11 short tons) in body mass. Using
11211-618: The upper Meuse roughly marked the western border of the Holy Roman Empire with the Kingdom of France , after Count Henry III of Bar had to receive the western part of the County of Bar ( Barrois mouvant ) as a French fief from the hands of King Philip IV . In 1408, a Burgundian army led by John the Fearless went to the aid of John III against the citizens of Liège, who were in open revolt. After
11322-483: The upper jaw, there were three types: the premaxillary teeth, maxillary teeth, and pterygoid teeth. On the lower jaw, only one type, the dentary teeth, were present. In each jaw row, from front to back, Mosasaurus had: two premaxillary teeth, twelve to sixteen maxillary teeth, and eight to sixteen pterygoid teeth on the upper jaw and fourteen to seventeen dentary teeth on the lower jaw. The teeth were largely consistent in size and shape with only minor differences throughout
11433-582: Was a type of derived mosasaur, or a latecoming member with advanced evolutionary traits such as a fully aquatic lifestyle. As such, it had a streamlined body, an elongated tail ending with a downturn supporting a two-lobed fin, and two pairs of flippers. While in the past derived mosasaurs were depicted as akin to giant flippered sea snakes , it is now understood that they were more similar in build to other large marine vertebrates such as ichthyosaurs, marine crocodylomorphs , and archaeocete whales through convergent evolution . The type species, M. hoffmannii ,
11544-613: Was confirmed in 2004, when the snout was found in the collections of the MNHN under the catalog number MNHN 958. The third species was described in 1881 by Edward Drinker Cope from a fragmentary fossil skeleton having been discovered in New Jersey , now cataloged as AMNH 1380. In his description, the Cope thought that it represented a giant species of Clidastes and named it Clidastes conodon . In 1966, Donald Baird and Gerard R. Case reidentified it as
11655-517: Was discovered in 1764 in a chalk quarry near Maastricht in the Netherlands in the form of a skull, which was initially identified as a whale . This specimen, cataloged as TM 7424, is now on display at the Teylers Museum in Haarlem . Later around 1780, the quarry produced a second skull that caught the attention of the physician Johann Leonard Hoffmann , who thought it was a crocodile . He contacted
11766-475: Was first described in 1834 by Richard Harlan based on a snout fragment found along the river's Big Bend , in South Dakota . In reference to its discovery made in the river, he coined the specific epithet and initially identified it as a species of Ichthyosaurus but later as an amphibian named Batrachiosaurus . The rest of the skull had been discovered earlier by a fur-trapper, and it eventually came under
11877-439: Was not stated whether they applied Russell's 1967 ratio, although Gayford et al. (2024) suggested it was likely. M. missouriensis and M. lemonnieri are smaller than M. hoffmannii but are known from more complete fossils. Based on measurements of various Belgian skeletons, Dollo estimated M. lemonnieri grew to around 7 to 10 meters (23 to 33 ft) in length. He also measured the dimensions of IRSNB 3119 and recorded that
11988-509: Was recently rebaptized into "Canal de la Meuse". The waterway can be used by the smallest barges that are still in use commercially almost 40 m (131 ft) long and just over 5 metres (16 ft) wide. Just upstream of the town of Commercy, the Canal de la Meuse connects with the Marne–Rhine Canal by means of a short diversion canal. The Cretaceous sea reptile Mosasaur is named after
12099-571: Was revolutionary at the time and helped support the then-developing ideas of extinction . Cuvier did not designate a scientific name for the animal; this was done by William Daniel Conybeare in 1822 when he named it Mosasaurus in reference to its origin in fossil deposits near the Meuse River. The exact affinities of Mosasaurus as a squamate remain controversial, and scientists continue to debate whether its closest living relatives are monitor lizards or snakes . The largest species, M. hoffmannii ,
12210-492: Was revolutionary at the time, and in 1812 he proclaimed, "Above all, the precise determination of the famous animal from Maastricht seems to us as important for the theory of zoological laws, as for the history of the globe." In a 1822 work by James Parkinson , William Daniel Conybeare coined the genus Mosasaurus from the Latin Mosa " Meuse " and the Ancient Greek σαῦρος ( saûros , "lizard"), all literally meaning "lizard of
12321-415: Was the landscape of Meuse by Joachim Patinir . He was likely the uncle of Henri Blès , who is sometimes defined as a Mosan landscape painter active during the second third of the 16th century (i.e. second generation of landscape painters). The main tributaries of the Meuse are listed below in downstream-upstream order, with the town where the tributary meets the river: The mean annual discharge rate of
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