title: Fascist Italy (1922–1943)

text: After Germany annexed Czechoslovakia, Mussolini turned his attention to Albania. On 7 April 1939, Italy invaded the country and after a short campaign, Albania was occupied, turned into a protectorate and its parliament crowned Victor Emmanuel III King of Albania. The historical justification for the annexation of Albania laid in the ancient history of the Roman Empire in which the region of Albania had been an early conquest for the Romans, even before Northern Italy had been taken by Roman forces. However, by the time of annexation little connection to Italy remained amongst Albanians. Albania was very closely tied to Italy even before the Italian invasion. Italy had built up heavy influence over Albania through the Treaties of Tirana, which gave Italy concessions over the Albanian economy and army. The occupation was not appreciated by King Emmanuel III, who feared that it had isolated Italy even further than its war against Ethiopia.


title: Fascist Italy (1922–1943)

text: In 1936 in Spain, the Fascist regime made its most significant pre-war military intervention. The Spanish Republic was divided in the Spanish Civil War between the anticlerical socialist Republicans and the Church-supporting nationalists led by Francisco Franco under fascist Falange movement. Italy sent aircraft, weapons and a total of over 60,000 troops to aid the Spanish nationalists. The war helped train the Italian military for war and improve relations with the Roman Catholic Church. It was a success that secured Italy's naval access in and out of the Mediterranean Sea to the Atlantic Ocean and its ability to pursue its policy of "Mare Nostrum" without fear of opposition by Spain. The other major foreign contributor to the Spanish Civil War was Germany. This was the first time that Italian and German forces fought together since the Franco-Prussian War in the 1870s. During the 1930s, Italy built many large battleships and other warships to solidify Italy's hold on the Mediterranean Sea.


title: Fascist Italy (1922–1943)

text: When the Nazi Party attained power in Germany in 1933, Mussolini and the Fascist regime in public showed approval of Hitler's regime, with Mussolini saying: "The victory of Hitler is our victory". The Fascist regime also spoke of creating an alliance with the new regime in Germany. In private, Mussolini and the Italian Fascists showed disapproval of the National Socialist government and Mussolini had a disapproving view of Hitler despite ideological similarities. The Fascists distrusted Hitler's Pan-German ideas which they saw as a threat to territories in Italy that previously had been part of the Austrian Empire. Although other National Socialists disapproved of Mussolini and Fascist Italy, Hitler had long idolized Mussolini's oratorical and visual persona and adopted much of the symbolism of the Fascists into the National Socialist Party, such as the Roman, straight-armed salute, dramatic oratory, the use of uniformed paramilitaries for political violence and the use of mass rallies to demonstrate the power of the movement. In 1922, Hitler tried to ask for Mussolini's guidance on how to organize his own version of the "March on Rome" which would be a "March on Berlin" (which came into being as the failed Beer Hall Putsch in 1923). Mussolini did not respond to Hitler's requests as he did not have much interest in Hitler's movement and regarded Hitler to be somewhat crazy. Mussolini did attempt to read "Mein Kampf" to find out what Hitler's National Socialist movement was, but was immediately disappointed, saying that "Mein Kampf" was "a boring tome that I have never been able to read" and remarked that Hitler's beliefs were "little more than commonplace clichés". While Mussolini like Hitler believed in the cultural and moral superiority of whites over colored peoples, he opposed Hitler's antisemitism. A number of Fascists were Jewish, including Mussolini's mistress Margherita Sarfatti, who was the director of Fascist art and propaganda, and there was little support amongst Italians for antisemitism. Mussolini also did not evaluate race as being a precursor of superiority, but rather culture.


title: Fascist Italy (1922–1943)

text: Hitler and the National Socialists continued to try to woo Mussolini to their cause and eventually Mussolini gave financial assistance to the Nazi Party and allowed National Socialist paramilitaries to train in Italy in the belief that despite differences, a nationalist government in Germany could be beneficial to Italy. As suspicion of the Germans increased after 1933, Mussolini sought to ensure that Germany would not become the dominant nationalist state in Europe. To do this, Mussolini opposed German efforts to annex Austria after the assassination of fascist Austrian President Engelbert Dollfuss in 1934 and promised the Austrians military support if Germany were to interfere. This promise helped save Austria from annexation in 1934.


title: Fascist Italy (1922–1943)

text: Public appearances and propaganda constantly portrayed the closeness of Mussolini and Hitler and the similarities between Italian Fascism and German National Socialism. While both ideologies had significant similarities, the two factions were suspicious of each other and both leaders were in competition for world influence. Hitler and Mussolini first met in June 1934, as the issue of Austrian independence was in crisis. In private after the visit in 1934, Mussolini said that Hitler was just "a silly little monkey".


title: Fascist Italy (1922–1943)

text: After Italy became isolated in 1936, the government had little choice but to work with Germany to regain a stable bargaining position in international affairs and reluctantly abandoned its support of Austrian independence from Germany. In September 1937, Mussolini visited Germany in order to build closer ties with his German counterpart. On 28 October 1937, Mussolini declared Italy's support of Germany regaining its colonies lost in World War I, declaring: "A great people such as the German people must regain the place which is due to it, and which it used to have beneath the sun of Africa".


title: Fascist Italy (1922–1943)

text: With no significant opposition from Italy, Hitler proceeded with the "Anschluss", the annexation of Austria in 1938. Germany later claimed the Sudetenland, a province of Czechoslovakia inhabited mostly by Germans. Mussolini felt he had little choice but to help Germany to avoid isolation. With the annexation of Austria by Germany in 1938, the Fascist regime began to be concerned about the majority ethnic German population in South Tyrol and whether they would want to join a Greater Germany. The Fascists were also concerned about whether Italy should follow National Socialist antisemitic policies in order to gain favor from those National Socialists who had mixed feelings about Italy as an ally. In 1938, Mussolini pressured fellow Fascist members to support the enacting of antisemitic policies, but this was not well taken as a number of Fascists were Jewish and antisemitism was not an active political concept in Italy. Nevertheless, Mussolini forced through antisemitic legislation even while his own son-in-law and prominent Fascist Count Galeazzo Ciano personally condemned such laws. In turn for enacting the extremely unpopular antisemitic laws, Mussolini and the Fascist government demanded a concession from Hitler and the National Socialists. In 1939, the Fascists demanded from Hitler that his government willingly accept the Italian government's plan to have all Germans in South Tyrol either leave Italy or be forced to accept Italianization. Hitler agreed and thus the threat to Italy from the South Tyrol Germans was neutralized.


title: Fascist Italy (1922–1943)

text: As war approached in 1939, the Fascist regime stepped up an aggressive press campaign against France claiming that Italian people were suffering in France. This was important to the alliance as both regimes mutually had claims on France, Germany on German-populated Alsace-Lorraine and Italy on Italian-populated Corsica, Nizza and Savoia. In May 1939, a formal alliance was organized. The alliance was known as the Pact of Steel, which obliged Italy to fight alongside Germany if war broke out against Germany. Mussolini felt obliged to sign the pact in spite of his own concerns that Italy could not fight a war in the near future. This obligation grew from his promises to Italians that he would build an empire for them and from his personal desire to not allow Hitler to become the dominant leader in Europe. Mussolini was repulsed by the Molotov–Ribbentrop Pact agreement where Germany and the Soviet Union agreed to partition the Second Polish Republic into German and Soviet zones for an impending invasion. The Fascist government saw this as a betrayal of the Anti-Comintern Pact, but decided to remain officially silent.


title: Fascist Italy (1922–1943)

text: Italy's military and logistical resources were stretched by successful pre-WWII military interventions in Spain, Ethiopia, Libya, and Albania and were not ready for a long conflict. Nevertheless, Mussolini went to war to further the imperial ambitions of the Fascist regime, which aspired to restore the Roman Empire in the Mediterranean (the "Mare Nostrum").


title: Fascist Italy (1922–1943)

text: Italy joined the war as one of the Axis Powers in 1940, entering after it appeared France was likely to lose to Germany. The Italian invasion of France was brief as the French Third Republic surrendered shortly afterward. Italy readied to fight against the British Empire in Africa and the Middle East, known as the "parallel war", while expecting a similar collapse of British forces in the European theatre. The Italians bombed Mandatory Palestine, invaded Egypt and occupied British Somaliland with initial success. The Italian military machine showed weakness during the 1940 Greco-Italian War, a war of aggression Italy launched unprovoked, but where the Italian army found little progress. German help during the Battle of Greece would eventually bail the Italians out, and their grander ambitions were partially met by late 1942 with Italian influence extended throughout the Mediterranean. Most of Greece was occupied by Italy; Italians administered the French territories of Corsica and Tunisia following Vichy France's collapse and occupation by German forces; and a puppet regime was installed in Croatia following the German-Italian Invasion of Yugoslavia. Albania, Ljubljana, coastal Dalmatia, and Montenegro had been directly annexed by the Italian state. Italo-German forces had also achieved victories against insurgents in Yugoslavia, and had occupied parts of British-held Egypt on their push to El-Alamein after their victory at Gazala.


title: Fascist Italy (1922–1943)

text: However, Italy's conquests were always heavily contested, both by various insurgencies (most prominently the Greek resistance and Yugoslav partisans) and Allied military forces, which waged the Battle of the Mediterranean throughout and beyond Italy's participation. German and Japanese actions in 1941 led to the entry of the Soviet Union and United States, respectively, into the war, thus ruining the Italian plan of forcing Britain to agree to a negotiated peace settlement. Ultimately the Italian empire collapsed after disastrous defeats in the Eastern European and North African campaigns. In July 1943, following the Allied invasion of Sicily, Mussolini was arrested by order of King Victor Emmanuel III, provoking the Italian Civil War. Italy's military outside of the Italian peninsula collapsed, its occupied and annexed territories falling under German control. Italy capitulated to the Allies on 3 September 1943.


title: Fascist Italy (1922–1943)

text: The northern half of the country was occupied by the Germans with the cooperation of Italian fascists, and became the Italian Social Republic, a collaborationist puppet state that recruited more than 500,000 soldiers for the Axis cause. The south was officially controlled by monarchist forces, which fought for the Allied cause as the Italian Co-Belligerent Army (at its height numbering more than 50,000 men), as well as around 350,000 Italian resistance movement partisans (mostly former Royal Italian Army soldiers) of disparate political ideologies that operated all over Italy. On 28 April 1945, Benito Mussolini was executed by Italian partisans, two days before Adolf Hitler's suicide.


title: Fascist Italy (1922–1943)

text: Most of the historiographical controversy centers on sharply conflicting interpretations of Fascism and the Mussolini regime. The 1920s writers on the left, following the lead of communist theorist Antonio Gramsci (1891–1937), stressed that Fascism was a form of capitalism. The Fascist regime controlled the writing and teaching of history through the central "Giunta Centrale per gli Studi Storici" and control of access to the archives and sponsored historians and scholars who were favorable toward it such as philosopher Giovanni Gentile and historians Gioacchino Volpe and Francesco Salata. In October 1932, it sponsored a large Exhibition of the Fascist Revolution, featuring its favored modernist art and asserting its own claims to express the spirit of Roman glory.


title: Fascist Italy (1922–1943)

text: After the war, most historiography was intensely hostile to Mussolini, emphasizing the theme of Fascism and totalitarianism. An exception was historian Renzo De Felice (1929–1996), whose Mussolini's Biography, four volumes and 6,000 pages long (1965–1997) remains the most exhaustive examination of public and private documents about Italian Fascism and serves as a basic resource for all scholars. De Felice argued that Mussolini was a revolutionary modernizer in domestic issues, but a pragmatist in foreign policy who continued the Realpolitik policies of liberal Italy (1861–1922). In the 1990s, a cultural turn began with studies that examined the issue of popular reception and acceptance of Fascism using the perspectives of "aestheticization of politics" and "sacralisation of politics". By the 21st century, the old "anti-Fascist" postwar consensus was under attack from a group of revisionist scholars who have presented a more favorable and nationalistic assessment of Mussolini's role, both at home and abroad. Controversy rages as there is no consensus among scholars using competing interpretations based on revisionist, anti-Fascist, intentionalist or culturalist models of history.


title: Styx

text: In Greek mythology, Styx (; ) is a river that forms the boundary between Earth (Gaia) and the Underworld. The rivers Acheron, Cocytus, Lethe, Phlegethon, and Styx all converge at the centre of the underworld on a great marsh, which sometimes is also called the Styx. According to Herodotus, the river Styx originates near Pheneus. Styx is also known as the goddess of the river, the source of its miraculous powers.


title: Styx

text: The deities of the Greek pantheon swore all their oaths upon the river Styx because, according to Greek mythology, during the Titanomachy, Styx, the goddess of the river, sided with Zeus. After the war, Zeus declared that every oath must be sworn upon her. Zeus swore to give Semele whatever she wanted and was then obliged to follow through when he realized to his horror that her request would lead to her death. Helios similarly promised his son Phaëton whatever he desired, also resulting in the boy's death. Myths related to such early deities did not survive long enough to be included in historic records, but tantalizing references exist among those that have been discovered.


title: Styx

text: According to some versions, the river Styx had miraculous powers which could make someone who bathed in the waters invulnerable. According to one tradition, Achilles' mother dipped him in the river during his childhood and he thus acquired invulnerability, with the exception of the heel by which his mother held him. Achilles was struck and killed during the Trojan War by an arrow shot into his heel by Paris. This is the source of the expression "Achilles' heel", a metaphor for a vulnerable spot.


title: Styx

text: Styx was primarily a feature in the afterworld of classical Greek mythology. The ferryman Charon is sometimes described as having transported the souls of the newly dead across this river into the Underworld, although in many sources it is instead the Acheron which Charon crosses and which is at the entrance of the Underworld. The further down the Styx people were carried, the longer and/or more severe would be their punishment. Dante, putting Charon and the Acheron at the entrance to Hell, put Phlegyas as ferryman over the Styx and made it the fifth circle of Hell, where the wrathful and sullen are punished by being drowned in the muddy waters for eternity, with the wrathful fighting each other.


title: Styx

text: Most historical accounts, including Pausanias (10.28) and later Dante's "Inferno" (3.78), associate Charon with the river Acheron. Ancient Greek literary sources – such as Pindar, Aeschylus, Euripides, Plato, and Callimachus – also place Charon on the Acheron. Roman poets, including Propertius, Ovid, and Statius, name the river as the Styx, perhaps following the geography of Virgil's underworld in the "Aeneid," where Charon is associated with both rivers.


title: Styx

text: In ancient times some believed that a coin (Charon's obol) placed in the mouth of a dead person would pay the toll for the ferry across the river to the entrance of the Underworld. It was said that if someone could not pay the fee, they would never be able to cross the river. The ritual was performed by the relatives of the dead. According to the myth Narcissus is still admiring himself in the Underworld, looking at the waters of the Styx.


title: Styx

text: The variant spelling Stix was sometimes used in translations of Classical Greek before the 20th century. By metonymy, the adjective "stygian" () came to refer to anything dark, dismal, and murky.


title: Styx

text: Styx was the name of an Oceanid nymph, one of the three thousand daughters of the Titans Oceanus and his sister-wife Tethys, and the goddess of the River Styx. Others make her one of the progenies of the primordial gods, Nyx (Night) and Erebus (Darkness). In classical myths, Styx's husband was Pallas and she gave birth to Zelus, Nike, Kratos, and Bia (and sometimes Eos and Selene). Another source, adds Scylla, the Fontes (Fountains), and the Lacus (Lakes) as their children. Styx also mothered Echidna by Peiras and even Persephone by Zeus in one account.


title: Styx

text: In these myths, Styx supported Zeus in the Titanomachy, where she was said to be the first to rush to his aid. For this reason, her name was given the honour of being a binding oath for the deities. Knowledge of whether this was the original reason for the tradition did not survive into historical records following the religious transition that led to the pantheon of the classical era.


title: Styx

text: On 2 July 2013, "Styx" officially became the name of one of Pluto's moons. The other moons of Pluto (Charon, Nix, Hydra, and Kerberos) also have names from Greco-Roman mythology related to the underworld.


title: Glam rock

text: Glam rock is a style of rock music that developed in the United Kingdom in the early 1970s and was performed by musicians who wore outrageous costumes, makeup, and hairstyles, particularly platform shoes and glitter. Glam artists drew on diverse sources across music and throwaway pop culture, ranging from bubblegum pop and 1950s rock and roll to cabaret, science fiction, and complex art rock. The flamboyant clothing and visual styles of performers were often camp or androgynous, and have been described as playing with other gender roles. Glitter rock was a more extreme version of glam rock.


title: Glam rock

text: The UK charts were inundated with glam rock acts from 1971 to 1975. The March 1971 appearance of T. Rex frontman Marc Bolan on the BBC's music show "Top of the Pops", wearing glitter and satins, is often cited as the beginning of the movement. Other British glam rock artists included David Bowie, Mott the Hoople, Sweet, Slade, Mud, Roxy Music and Gary Glitter. Those not central to the genre, artists such as Elton John, Rod Stewart and Freddie Mercury of Queen, also adopted glam styles. In the United States, the scene was much less prevalent, with Alice Cooper and Lou Reed the only American artists to score a hit in the UK. Other American glam artists include New York Dolls, Sparks, Suzi Quatro, Iggy Pop and Jobriath. It declined after the mid-1970s, but influenced other musical genres including punk rock, glam metal, New Romantic, death rock and gothic rock.


title: Glam rock

text: Glam rock can be seen as a fashion as well as musical subgenre. Glam artists rejected the revolutionary rhetoric of the late 1960s rock scene, instead glorifying decadence, superficiality, and the simple structures of earlier pop music. In response to these characteristics, scholars such as I.Taylor and D. Wall characterised glam rock as "offensive, commercial, and cultural emasculation".


title: Glam rock

text: Artists drew on such musical influences as bubblegum pop, the brash guitar riffs of hard rock, stomping rhythms, and 1950s rock and roll, filtering them through the recording innovations of the late 1960s. Ultimately, it became very diverse, varying between the simple rock and roll revivalism of figures like Alvin Stardust to the complex art pop of Roxy Music. In its beginning, however, it was a youth-orientated reaction to the creeping dominance of progressive rock and concept albums – what "Bomp!" called the "overall denim dullness" of "a deadly boring, prematurely matured music scene".


title: Glam rock

text: Visually, it was a mesh of various styles, ranging from 1930s Hollywood glamour, through 1950s pin-up sex appeal, pre-war cabaret theatrics, Victorian literary and symbolist styles, science fiction, to ancient and occult mysticism and mythology; manifesting itself in outrageous clothes, makeup, hairstyles, and platform-soled boots. Glam rock is most noted for its sexual and gender ambiguity and representations of androgyny, beside extensive use of theatrics.


title: Glam rock

text: It was prefigured by the flamboyant English composer Noël Coward, especially his 1931 song "Mad Dogs and Englishmen", with music writer Daryl Easlea stating, "Noël Coward's influence on people like Bowie, Roxy Music and Cockney Rebel was absolutely immense. It suggested style, artifice and surface were equally as important as depth and substance. "Time" magazine noted Coward's 'sense of personal style, a combination of cheek and chic, pose and poise'. It reads like a glam manifesto." Showmanship and gender identity manipulation acts included the Cockettes and Alice Cooper, the latter of which combined glam with shock rock.


title: Glam rock

text: Glam rock emerged from the English psychedelic and art rock scenes of the late 1960s and can be seen as both an extension of, and a reaction against, those trends. Its origins are associated with Marc Bolan, who had renamed his acoustic duo T. Rex and taken up electric instruments by the end of the 1960s. Bolan was, in the words of music critic Ken Barnes, "the man who started it all". Often cited as the moment of inception is Bolan's appearance on the BBC music show "Top of the Pops" in March 1971 wearing glitter and satins, to perform what would be his second UK Top 10 hit (and first UK Number 1 hit), "Hot Love". "The Independent" states that Bolan's appearance on "Top of the Pops" "permitted a generation of teeny-boppers to begin playing with the idea of androgyny". T. Rex's 1971 album "Electric Warrior" received critical acclaim as a pioneering glam rock album. In 1973, a few months after the release of the album "Tanx", Bolan captured the front cover of "Melody Maker" magazine with the declaration "Glam rock is dead!"


title: Glam rock

text: From late 1971, already a minor star, David Bowie developed his Ziggy Stardust persona, incorporating elements of professional makeup, mime and performance into his act. Bowie, in a 1972 interview in which he noted that other artists described as glam rock were doing different work, said "I think glam rock is a lovely way to categorize me and it's even nicer to be one of the leaders of it". Bolan and Bowie were soon followed in the style by acts including Roxy Music, Sweet, Slade, Mott the Hoople, Mud and Alvin Stardust. The popularity of glam rock in the UK was such that three glam rock bands had major UK Christmas hit singles; "Merry Xmas Everybody" by Slade, "I Wish It Could Be Christmas Everyday" by Wizzard and "Lonely This Christmas" by Mud, all of which have remained hugely popular. Glam was not only a highly successful trend in UK popular music, it became dominant in other aspects of British popular culture during the 1970s.


title: Glam rock

text: A heavier variant of glam rock, emphasising guitar riff centric songs, driving rhythms and live performance with audience participation, were represented by bands like Slade and Mott the Hoople, with later followers such as Def Leppard, Cheap Trick, Poison, Kiss, and Quiet Riot, some of which either covered Slade compositions (such as "Cum On Feel the Noize" and "Mama Weer All Crazee Now") or composed new songs based on Slade templates. While highly successful in the single charts in the UK (Slade for example had six number one singles), very few of these musicians were able to make a serious impact in the US; David Bowie was the major exception, becoming an international superstar and prompting the adoption of glam styles among acts like Lou Reed, Iggy Pop, New York Dolls and Jobriath, often known as "glitter rock" and with a darker lyrical content than their British counterparts.


title: Glam rock

text: In the UK, the term glitter rock was most often used to refer to the extreme version of glam pursued by Gary Glitter and the independent band with whom he often performed known as the Glitter Band. The Glitter Band and Gary Glitter had between them eighteen top ten singles in the UK between 1972 and 1975. A second wave of glam rock acts, including Suzi Quatro, Roy Wood's Wizzard and Sparks, had hits on the British single charts in 1973 and 1974. Quatro directly inspired the pioneering Los Angeles based all-girl group The Runaways. Existing acts, some not usually considered central to the genre, also adopted glam styles, including Rod Stewart, Elton John, Queen and, for a time, The Rolling Stones. After seeing Marc Bolan wearing Zandra Rhodes-designed outfits, Freddie Mercury enlisted Rhodes to design costumes for the next Queen tour in 1974. Punk rock, often seen as a reaction to the artifice of glam rock, but using some elements of the genre, including makeup and involving cover versions of glam rock records, helped end the fashion for glam from about 1976.


title: Glam rock

text: While glam rock was exclusively a British cultural phenomenon, with Steven Wells in "The Guardian" writing "Americans only got glam second hand via the posh Bowie version", covers of British glam rock classics are now piped-muzak staples at US sporting events. Glam rock was a background influence for Richard O'Brien, writer of the 1973 London musical "The Rocky Horror Show". Although glam rock went into a steep decline in popularity in the UK in the second half of the 1970s, it had a direct influence on acts that rose to prominence later, including Kiss and American glam metal acts like Quiet Riot, W.A.S.P., Twisted Sister, Bon Jovi and Mötley Crüe.


title: Glam rock

text: New Romantic acts in the UK such as Adam and the Ants and A Flock of Seagulls extended glam, and its androgyny and sexual politics were picked up by acts including Culture Club, Bronski Beat and Frankie Goes to Hollywood. Gothic rock was largely informed by the makeup, clothes, theatricality and sound of glam, and punk rock adopted some of the performance and persona-creating tendencies of glam, as well as the genre's emphasis on pop-art qualities and simple but powerful instrumentation.


title: Glam rock

text: Glam rock has been influential around the world. In Japan in the 1980s, visual kei was strongly influenced by glam rock aesthetics. Glam rock has since enjoyed continued influence and sporadic modest revivals in R&B crossover act Prince, bands such as Marilyn Manson, Suede, Placebo, Chainsaw Kittens, Spacehog and the Darkness, and has inspired pop artists such as Lady Gaga.


title: Richard Cottingham

text: Richard Francis Cottingham (born November 25, 1946) is an American serial killer and rapist who murdered at least 17 young women and girls in New York and New Jersey between 1967 and 1980. His confirmed killings include nine convictions and a further eight official confessions under non-prosecution agreements. He was nicknamed The Torso Killer and The Times Square Killer. In 2009, decades after being convicted of five murders in New Jersey and New York in 1981–1984, Cottingham told a journalist that he had committed at least 80 to 100 "perfect murders" of women in various regions of the United States. Four surviving victims testified against Cottingham; he was convicted in three of the abduction-rape survivor cases and acquitted in one. He has been incarcerated in New Jersey state prisons since 1980, and has accumulated multiple life sentences.


title: Richard Cottingham

text: Cottingham was born on November 25, 1946, in Mott Haven, Bronx, New York City, the first of four children (he has three sisters). In 1948, his family moved to Dumont, New Jersey, and in 1956 to River Vale, New Jersey, where he began his fascination with bondage pornography. In 1964, Cottingham graduated from Pascack Valley High School in Hillsdale, New Jersey.


title: Richard Cottingham

text: In 1964, after graduating from high school, Cottingham began working for Metropolitan Life, where his father was a vice-president, at the firm's headquarters office on Madison Avenue in Manhattan in New York City. He started in the mail room and eventually became a mainframe computer operator after taking computer courses. In October 1966 he became a computer operator for Blue Cross Blue Shield Association in New York, where he worked until his 1980 arrest. At Blue Cross, Cottingham worked in an office with fugitive serial killer Rodney Alcala, the "Dating Game Killer", who was living in New York in 1969 under the alias "John Berger". Neither man has claimed to have been aware of the other, nor is there any evidence they were familiar with one another prior to their respective arrests.


title: Richard Cottingham

text: On May 3, 1970, Cottingham was married at Our Lady of Lourdes Church in Queens Village, New York. He and his wife had three children, two boys and a girl. In April 1978, his wife filed for divorce on the grounds of "abandonment" and "mental cruelty" (refusing to have sex with her after the birth of their third child, staying out until early morning, and leaving her with insufficient household funds). His wife withdrew the petition upon his arrest in May 1980, until after he was convicted in the first trial in New Jersey, then refiled the petition and completed the divorce proceeding.


title: Richard Cottingham

text: Cottingham was arrested on several lesser charges throughout his killing spree; the police were not aware of his murders at the time, nor were they aware there was an active serial killer at large in the New York-New Jersey area.


title: Richard Cottingham

text: On October 3, 1969, Cottingham was charged and convicted of drunk driving in New York City, and sentenced to a $50 fine.


title: Richard Cottingham

text: On August 21, 1972, Cottingham was charged and convicted of shoplifting at Stern's Department Store in Paramus, New Jersey and was sentenced to pay a $50 fine or ten days in jail. He paid the fine.


title: Richard Cottingham

text: On September 4, 1973, Cottingham was arrested in New York City for robbery, oral sodomy and sex abuse on the complaint of a prostitute and her pimp. Neither complainant appeared in further proceedings, however, and the case was dismissed.


title: Richard Cottingham

text: On March 12, 1974, Cottingham was arrested in New York City for robbery and unlawful imprisonment on the complaint of a prostitute. Once again, the victim did not appear in further proceedings, and the case was dismissed.


title: Richard Cottingham

text: Cottingham committed his first known murder when he was 20 years old. On October 28, 1967, he strangled Nancy Schiava Vogel, a 29-year-old married mother of two, in Little Ferry, New Jersey. Her nude body, hands bound in front of her, was found on October 31, under a blanket behind the passenger seat of her car parked in Ridgefield Park, New Jersey. She had last been seen three days earlier, when she left home stating she was going to play bingo with friends at a local church. The murder remained unsolved until Cottingham confessed and pleaded guilty to it in August 2010.


title: Richard Cottingham

text: On February 15, 1968, Cottingham killed his second known victim. After failing to return home from a trip to buy shoes at the Green Acres Mall in Valley Stream, New York, 23-year old Diane Cusick was found raped, beaten, and strangled to death in the back seat of her car, parked near the mall. Cottingham was not charged with this murder until June 2022, when he was implicated by DNA.


title: Richard Cottingham

text: Starting in 2014, Cottingham confidentially admitted to Detective Robert Anzilotti of the Bergen County Prosecutor's Office, New Jersey, the murders of three teenage females in 1968 and 1969:


title: Richard Cottingham

text: The Bergen County Prosecutor's Office (BCPO) "exceptionally closed" the three cold case murders with agreement from the victims' families, but for several years kept this secret from the public to keep Cottingham talking about other cases. In December 2019, forensic historian and author Peter Vronsky, on the eve of publishing the revelation in his second edition of "", made it public, with BCPO cooperation, in a community meeting in Midland Park, the site of Jacalyn Harp's murder. Anzilotti and BCPO subsequently confirmed the "exceptional closures" of the three schoolgirl murders from 1968 to 1969.


title: Richard Cottingham

text: In August 2022, with a non-prosecution agreement, officials in Rockland County, New York, accepted Cottingham's confession to the murder of 26-year-old Lorraine McGraw, who was beaten and killed March 1, 1970.


title: Richard Cottingham

text: In December 2022, Cottingham was convicted of Cusick's 1968 murder and, under a non-prosecution agreement, officially admitted to another four killings during 1972–1973 in Long Island, New York: Mary Beth Heinz, Laverne Moye, Sheila Heiman, and Maria Emerita Rosado Nieves.


title: Richard Cottingham

text: In April 2021, Cottingham confessed to the unsolved August 9–11, 1974, double-abduction, rape, and forcible drowning murders of Lorraine Marie Kelly, 16, and Mary Ann Pryor, 17, in Montvale, New Jersey, one of New Jersey's most notorious cold cases. The confession was extracted by Chief of Detectives Robert Anzilotti weeks before his retirement and was facilitated by the work of forensic historian Peter Vronsky and a Cottingham victim family member, Jennifer Weiss, the daughter of Deedeh Goodarzi, one of the New York torso killing victims. Vronsky and Weiss had been meeting with Cottingham in prison since the spring of 2017, counseling him to make the confession. Anzilotti had spent 15 years interviewing Cottingham, working toward the confession, which raised the total number of victims officially attributed to Cottingham to 11 at that time. He claims to have committed between 85 and 100 murders.


title: Richard Cottingham

text: Cottingham was convicted of five 1977–1980 murders in a series of three trials during 1981–1984, two in New Jersey and one in New York:


title: Richard Cottingham

text: On May 22, 1980, Cottingham picked up 18-year-old Leslie Ann O'Dell, who was soliciting on the corner of Manhattan's Lexington Avenue and 25th Street. At some point, she agreed to have sex with him for $44. Around dawn, they checked into the same Hasbrouck Heights Quality Inn where he had, eighteen days earlier, left Valerie Street's handcuffed body under a bed to be discovered by a housekeeper. Cottingham offered to give O'Dell a massage and she rolled onto her stomach. Straddling her back, he drew a knife and put it to her throat as he snapped a pair of handcuffs on her wrists.


title: Richard Cottingham

text: He began torturing her, nearly biting off one of her nipples. She later testified that he said, "You have to take it. The other girls did, you have to take it too. You're a whore and you have to be punished." O'Dell's muffled cries of pain became so loud that the motel staff, already spooked by the murder eighteen days earlier, called police and then rushed to the room demanding that Cottingham open the door. Cottingham was apprehended in the hallway by arriving police officers. When arrested, he had handcuffs, a leather gag, two slave collars, a switchblade knife, replica pistols and a stockpile of prescription pills.


title: Richard Cottingham

text: The charges listed in Cottingham's New Jersey indictment included kidnapping, attempted murder, aggravated assault, aggravated assault with deadly weapon, aggravated sexual assault while armed (rape), aggravated sexual assault while armed (sodomy), aggravated sexual assault while armed (fellatio), possession of a weapon, possession of controlled dangerous substances Secobarbital, Amobarbital and Diazepam.


title: Richard Cottingham

text: In April 1978, after his wife had initiated divorce proceedings, he kept a locked room in a basement apartment of the house in which they lived in Lodi, New Jersey. Following his 1980 arrest, police found, in the locked room and in the trunk of his car, personal effects which they traced to two of his victims.


title: Richard Cottingham

text: During the early 1980s, Cottingham was convicted of five murders – in two separate New Jersey murder trials in 1981 and 1982, and in a single trial in New York City in 1984 for three murders. He pleaded innocent and for decades insisted he was being "framed", until admitting in 2009 that he had actually perpetrated the murders he was accused of. Cottingham was apparently "forensically aware" and, in the 13-year period during which he is known to have committed at least 17 murders, in the pre-DNA era, only one fingerprint belonging to him was ever recovered, from the ratchet mechanism of handcuffs left behind on Valerie Street. A case based on his "signature pattern" was built against him, combined with the testimony of four surviving victims, as well as pieces of his victims' jewelry and other items found in his possession after his arrest.


title: Richard Cottingham

text: In 2010, he pleaded guilty to the 1967 murder of Nancy Vogel. During 2014–2017, in exchange for immunity from prosecution, he confessed to murdering three New Jersey schoolgirls in 1968–1969.


title: Richard Cottingham

text: In 2021, he pleaded guilty to the 1974 kidnapping, raping and drowning of Lorraine Marie Kelly and Mary Ann Pryor.


title: Richard Cottingham

text: In 2022, Cottingham officially confessed to the 1970 murder of Lorraine McGraw, under a non-prosecution agreement. His additional confession to a 1974 murder was discounted by Rockland County police.


title: Richard Cottingham

text: In 2022, he was arraigned from his hospital bed for the 1968 murder of Diane Cusick in Long Island, New York. The link was found through DNA evidence; authorities believed it to be, thus far, the oldest criminal case to be prosecuted by DNA evidence. He pleaded guilty in a December court appearance, and also officially admitted killing four other women in Long Island during 1972–1973: Mary Beth Heinz, Laverne Moye, Sheila Heiman, and Maria Emerita Rosado Nieves.


title: Richard Cottingham

text: Cottingham's case has been discussed in several books and documentaries on serial killers. Two focused entirely on him: Rod Leith's "The Prostitute Murders: The People vs. Richard Cottingham" (Lyle Stuart Inc., 1983) and "" (Netflix, 2021).


title: Mosasaurus

text: Mosasaurus (; "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 earliest fossils of "Mosasaurus" known to science were found as skulls in a chalk quarry near the Dutch city of Maastricht in the late 18th century, which were initially thought to have been the bones of crocodiles or whales. One skull discovered around 1780, and which was seized by France during the French Revolutionary Wars for its scientific value, 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 was revolutionary at the time and helped support the then-developing ideas of extinction. Cuvier did not designate a scientific name for the new animal, and 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.


title: Mosasaurus

text: Traditional interpretations have estimated the maximum length of the largest species, "M. hoffmannii", to be up to , making it one of the largest mosasaurs, although some scientists consider this an overestimation with recent estimates suggesting a length closer to . The skull of "Mosasaurus" was equipped with robust jaws capable of swinging back and forth and strong muscles capable of powerful bites using dozens of large teeth adapted for cutting prey. Its four limbs were shaped into robust paddles to steer the animal underwater. Its tail was long and ended in a downward bend and a paddle-like fluke. "Mosasaurus" was a predator possessing excellent vision to compensate for its poor sense of smell, and a high metabolic rate suggesting it was endothermic ("warm-blooded"), an adaptation only found in mosasaurs among squamates. 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 a historically problematic classification. As a result, more than fifty different 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 in a future study.


title: Mosasaurus

text: Fossil evidence suggests "Mosasaurus" inhabited much of the Atlantic Ocean and the seaways adjacent to it. "Mosasaurus" fossils have been found in places as diverse as 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 climates. "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 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 preythough they were able to coexist in 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, similarly seen in modern crocodiles today.


title: Mosasaurus

text: The first "Mosasaurus" fossil known to science 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. 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 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 1798 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 the fossil into cultural fame, but historians agree that the narrative was exaggerated.


title: Mosasaurus

text: 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 1808, Camper's son Adriaan Gilles Camper and Georges Cuvier concluded that the fossil, which by then was nicknamed the "great animal of Maastricht", belonged to a marine lizard with affinities to monitor lizards, but otherwise unlike any modern animal. 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 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." William Daniel Conybeare coined the genus "Mosasaurus" in 1822, and Gideon Mantell added the specific epithet "hoffmannii" in 1829. Cuvier later designated the second skull as the new species' holotype (defining example).


title: Mosasaurus

text: In 1804, the Lewis and Clark Expedition discovered a now-lost fossil skeleton alongside the Missouri River, which was identified as a 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.


title: Mosasaurus

text: The type specimen of "M. missouriensis" was first described in 1834 by Richard Harlan based on a snout fragment found along the river's Big Bend. He coined the specific epithet and initially identified it as a species of "Ichthyosaurus" but later as an amphibian. The rest of the skull had been discovered earlier by a fur-trapper, and it eventually came under the possession of prince Maximilian of Weid-Neuwied between 1832 and 1834. The fossil 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 was confirmed in 2004.


title: Mosasaurus

text: The third species was described in 1881 from fragmentary fossils in New Jersey by Edward Drinker Cope, who thought it was a giant species of "Clidastes" and named it "Clidastes conodon". In 1966, it was reidentified as a species of "Mosasaurus".


title: Mosasaurus

text: 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, but Cuvier rejected the idea of another "Mosasaurus" species. This species was re-introduced to science and formally described in 1889 by Louis Dollo based on a skull recovered from a phosphate quarry in Belgium. Further mining of 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 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 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.


title: Mosasaurus

text: "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 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 Ikejiri and Lucas in 2014. In 2004, Eric Mulder, Dirk Cornelissen, and Louis Verding suggested "M. lemonnieri" could be 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.


title: Mosasaurus

text: 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.


title: Mosasaurus

text: 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 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 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.


title: Mosasaurus

text: 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 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 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).


title: Mosasaurus

text: "Mosasaurus" 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.


title: Mosasaurus

text: The type species, "M. hoffmannii", is one of the largest mosasaurs 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 in length) to estimate a maximum length of . Using a smaller partial jaw (NHMM 009002) measuring and "reliably estimated at" when complete, Lingham-Soliar (1995) estimated a larger maximum length of 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 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 would have been up to or more than in length and weighed in body mass. Mentioning the Penza specimen, Gregory S. Paul estimated in his 2022 book, "The Princeton Field Guide to Mesozoic Sea Reptiles", a shorter maximum length for "M. hoffmannii" of and a body mass of .


title: Mosasaurus

text: 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 in length. It was not stated whether they applied Russell's 1967 ratio.


title: Mosasaurus

text: "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 in length. He also measured the dimensions of IRSNB 3119 and recorded that the skull constituted approximately one-eleventh of the whole body. Paul (2022) offered a larger maximum estimate for the species at in length and in body mass. Polcyn "et al." (2014) estimated that "M. missouriensis" may have measured up to in length. Street (2016) noted that large "M. missouriensis" individuals typically had skulls exceeding lengths of . A particular near-complete skeleton of "M. missouriensis" is reportedly measured at in total length with a skull approaching in length.; Paul (2022) estimated an individual of that size to weigh . Based on personal observations of various unpublished fossils from Morocco, Nathalie Bardet estimated that "M. beaugei" grew to a total length of , their skulls typically measuring around in length, with a body mass of around per Paul (2022). With a skull measuring around in length, "M. conodon" has been regarded as a small to medium-sized representative of the genus; Paul (2022) estimated its maximum length as being and body mass as being .


title: Mosasaurus

text: 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 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 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 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.


title: Mosasaurus

text: The palate, which consists of 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 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 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.


title: Mosasaurus

text: The features of teeth in "Mosasaurus" vary across species, but unifying characteristics include 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 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.


title: Mosasaurus

text: Like all mosasaurs, "Mosasaurus" had four types of teeth, classified based on the jaw bones they were located on. On 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 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 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.


title: Mosasaurus

text: 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 a "M. hoffmannii" maxillary tooth measured an average rate of deposition of odontoblasts, the cells responsible for the formation of dentin, at 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.


title: Mosasaurus

text: One of 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 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 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.


title: Mosasaurus

text: 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 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 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".


title: Mosasaurus

text: 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 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 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.


title: Mosasaurus

text: Because nomenclatural rules were not well-defined at the time, 19th century scientists did not give "Mosasaurus" 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 a future formal reassessment. Street & Caldwell (2017) was derived from Street's 2016 doctoral thesis, which contained a phylogenetic study prosposing 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 belonging to different genera and "M. beaugei" being a synonym of "M. hoffmannii".


title: Mosasaurus

text: 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 a tribe within the Mosasaurinae variously called Mosasaurini or Plotosaurini.


title: Mosasaurus

text: 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.


title: Mosasaurus

text: In 1997, Bell published 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 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.


title: Mosasaurus

text: Bell's study served as a precedent for later studies that mostly left the systematics of "Mosasaurus" unchanged, although some later studies have recovered 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 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 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.


title: Mosasaurus

text: 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 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.


title: Mosasaurus

text: The following cladogram on 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 the Mosasaurinae, with proposed new taxa and renamings in single quotations.


title: Mosasaurus

text: In 1995, Lingham-Soliar studied the head musculature of "M. hoffmannii". Because soft tissue like muscles do not easily fossilize, reconstruction of the musculature was largely based on the structure of the skull, muscle scarring on the skull, and the musculature in extant monitor lizards.


title: Mosasaurus

text: In modern lizards, the mechanical build of the skull is characterized by a four-pivot geometric structure in the cranium that allows flexible movement of the jaws, possibly to allow the animals to better position them and prevent prey escape when hunting. In contrast, the frontal and parietal bones, which in modern lizards connect to form a flexible pivot point, overlap in the skull of "M. hoffmannii". This creates a rigid three-pivot geometric cranial structure. These cranial structures are united by strong interlocking sutures formed to resist compression and shear forces caused by a downward thrust of the lower jaw muscles or an upward thrust of prey. This rigid but highly shock-absorbent structure of the cranium likely allowed a powerful bite force.


title: Mosasaurus

text: Like all mosasaurs, the lower jaws of "Mosasaurus" could swing forward and backward. In many mosasaurs like "Prognathodon" and "M. lemonnieri", this function mainly served to allow ratchet feeding, in which the pterygoid and jaws would "walk" captured prey into the mouth like a conveyor belt. But especially compared to those in "M. lemonnieri", the pterygoid teeth in "M. hoffmannii" are relatively small, which indicates ratchet feeding was relatively unimportant to its hunting and feeding. Rather, "M. hoffmannii" likely employed inertial feeding (in which the animal thrusts its head and neck backward to release a held prey item and immediately thrust the head and neck forward to close the jaws around the item) and used jaw adduction to assist in biting during prey seizure. The "magnus adductor" muscles, which attach to the lower jaws to the cranium and have a major role in biting function, are massive, indicating "M. hoffmannii" was capable of enormous bite forces. The long, narrow, and heavy nature of the lower jaws and attachment of tendons at the coronoid process would have allowed quick opening and closing of the mouth with little energy input underwater, which also contributed to the powerful bite force of "M. hoffmannii" and suggests it would not have needed the strong "magnus depressor" muscles (jaw-opening muscles) seen in some plesiosaurs.


title: Mosasaurus

text: "Mosasaurus" swam using its tail. The swimming style was likely sub-carangiform, which is exemplified today by mackerels. Its elongated paddle-like limbs functioned as hydrofoils for maneuvering the animal. The paddles' steering function was enabled by large muscle attachments from the outwards-facing side of the humerus to the radius and ulna and modified joints allowed an enhanced ability of rotating the flippers. The powerful forces resulting from utilization of the paddles may have sometimes resulted in bone damage, as evidenced by a "M. hoffmannii" ilium with significant separation of the bone's head from the rest of the bone likely caused by frequent shearing forces at the articulation joint.


title: Mosasaurus

text: The tissue structure of "Mosasaurus" bones suggests it had a metabolic rate much higher than modern squamates and its resting metabolic rate was between that of the leatherback sea turtle and that of ichthyosaurs and plesiosaurs. "Mosasaurus" was likely endothermic and maintained a constant body temperature independent of the external environment. Although there is no direct evidence specific to the genus, studies on the biochemistry of related mosasaur genera such as "Clidastes" suggests that endothermy was likely present in all mosasaurs. Such a trait is unique among squamates, the only known exception being the Argentine black and white tegu, which can maintain partial endothermy. This adaptation would have given several advantages to "Mosasaurus", including increased stamina when foraging across larger areas and pursuing prey. It may have also been a factor that allowed "Mosasaurus" to thrive in the colder climates of locations such as Antarctica.


title: Mosasaurus

text: "Mosasaurus" had relatively large eye sockets with large sclerotic rings occupying much of the sockets' diameter; the latter is correlated with eye size and suggests it had good vision. The eye sockets were located at the sides of the skull, which created a narrow field of binocular vision at around 28.5° but alternatively allowed excellent processing of a two-dimensional environment, such as the near-surface waters inhabited by "Mosasaurus".


title: Mosasaurus

text: Brain casts made from fossils of "Mosasaurus" show that the olfactory bulb and vomeronasal organ, which both control the function of smell, are poorly developed and lack some structures in "M. hoffmannii"; this indicates the species had a poor sense of smell. In "M. lemonnieri", these olfactory organs, although still small, are better developed and have some components lacking in "M. hoffmannii". The lack of a strong sense of smell suggests that olfaction was not particularly important in "Mosasaurus"; instead, other senses like vision may have been more useful.


title: Mosasaurus

text: Paleontologists generally agree that "Mosasaurus" was likely an active predator of a variety of marine animals. Fauna likely preyed upon by the genus include bony fish, sharks, cephalopods, birds, and marine reptiles such as other mosasaurs and turtles. It is unlikely "Mosasaurus" was a scavenger as it had a poor sense of smell. "Mosasaurus" was among the largest marine animals of its time, and with its large, robust cutting teeth, scientists believe larger members of the genus would have been able to handle virtually any animal. Lingham-Soliar (1995) suggested that "Mosasaurus" had a rather "savage" feeding behavior as demonstrated by large tooth marks on scutes of the giant sea turtle "Allopleuron hoffmanni" and fossils of re-healed fractured jaws in "M. hoffmannii". The species likely hunted near the ocean surface as an ambush predator, using its large two-dimensionally adapted eyes to more effectively spot and capture prey. Chemical and structural data in the fossils of "M. lemonnieri" and "M. conodon" suggests they may have also hunted in deeper waters.


title: Mosasaurus

text: Carbon isotope studies on fossils of multiple "M. hoffmannii" individuals have found extremely low values of δC, the lowest in all mosasaurs for the largest individuals. Mosasaurs with lower δC values tended to occupy higher trophic levels, and one factor for this was dietary: a diet of prey rich in lipids such as sea turtles and other large marine reptiles can lower δC values. "M. hoffmannii"'s low δC levels reinforces its likely position as an apex predator.


title: Mosasaurus

text: Currently, there is only one known example of a "Mosasaurus" preserved with stomach contents: a well-preserved partial skeleton of a small "M. missouriensis" dated about 75 million years old with dismembered and punctured remains of a long fish in its gut. This fish was much longer than the length of the mosasaur's skull, which measured in length, confirming that "M. missouriensis" consumed prey larger than its head by dismembering and consuming bits at a time. Due coexistence with other large mosasaurs like "Prognathodon", which specialized in robust prey, "M. missouriensis" likely specialized more on prey best consumed using cutting-adapted teeth in an example of niche partitioning.


title: Mosasaurus

text: "Mosasaurus" may have taught their offspring how to hunt, as supported by a fossil nautiloid "Argonautilus catarinae" with bite marks from two conspecific mosasaurs, one being from a juvenile and the other being from an adult. Analysis of the tooth marks by a 2014 study by Kauffman concluded that the mosasaurs were either "Mosasaurus" or "Platecarpus". The positioning of both bite marks are at the direction the nautiloid's head would have been facing, indicating it was incapable of escaping and was thus already sick or dead during the attacks; it is possible this phenomenon was from a parent mosasaur teaching its offspring about cephalopods as an alternate source of prey and how to hunt one. An alternate explanation postulates the bite marks as from one individual mosasaur that lightly bit the nautiloid at first, then proceeded to bite again with greater force. However, there are differences in tooth spacing between both bites which indicate different jaw sizes.


title: Mosasaurus

text: There is fossil evidence that "Mosasaurus" engaged in aggressive and lethal combat with others of its kind. One partial skeleton of "M. conodon" bears multiple cuts, breaks, and punctures on various bones, particularly in the rear portions of the skull and neck, and a tooth from another "M. conodon" piercing through the quadrate bone. No injuries on the fossil show signs of healing, suggesting that the mosasaur was killed by its attacker by a fatal blow in the skull. Likewise, an "M. missouriensis" skeleton has a tooth from another "M. missouriensis" embedded in the lower jaw underneath the eye. In this case, there were signs of healing around the wound, implying survival of the incident. Takuya Konishi suggested an alternative cause of this example being head-biting behavior during courtship as seen in modern lizards.


title: Mosasaurus

text: Attacks by another "Mosasaurus" are a possible cause of physical pathologies in other skulls, but they could have instead arisen from other incidents like attempted biting on hard turtle shells. In 2004, Lingham-Soliar observed that if these injuries were indeed the result of an intraspecific attack, then there is a pattern of them concentrating in the skull region. Modern crocodiles commonly attack each other by grappling an opponent's head using their jaws, and Lingham-Soliar hypothesized that "Mosasaurus" employed similar head-grappling behavior during intraspecific combat. Many of the fossils with injuries possibly attributable to intraspecific combat are of juvenile or sub-adult "Mosasaurus", leading to the possibility that attacks on smaller, weaker individuals may have been more common. However, the attacking mosasaurs of the "M. conodon" and "M. missouriensis" specimens were likely similar in size to the victims. In 2006, Schulp and colleagues speculated that "Mosasaurus" may have occasionally engaged in cannibalism as a result of intraspecific aggression.


title: Mosasaurus

text: There are some "M. hoffmannii" jaws with evidence of infectious diseases as a result of physical injuries. Two examples include IRSNB R25 and IRSNB R27, both having fractures and other pathologies in their dentaries. IRSNB R25 preserves a complete fracture near the sixth tooth socket. Extensive amounts of bony callus almost overgrowing the tooth socket are present around the fracture along with various osteolytic cavities, abscess canals, damages to the trigeminal nerve, and inflamed erosions signifying severe bacterial infection. There are two finely ulcerated scratches on the bone callus, which may have developed as part of the healing process. IRSNB R27 has two fractures: one had almost fully healed and the other is an open fracture with nearby teeth broken off as a result. The fracture is covered with a nonunion formation of bony callus with shallow scratch marks and a large pit connected to an abscess canal. Lingham-Soliar described this pit as resembling a tooth mark from a possible attacking mosasaur. Both specimens show signs of deep bacterial infection alongside the fractures; some bacteria may have spread to nearby damaged teeth and caused tooth decay, which may have entered deeper tissue from prior post-traumatic or secondary infections. The dentaries ahead of the fractures in both specimens are in good condition, suggesting that the arteries and trigeminal nerves had not been damaged; if they were, those areas would have necrotized due to lack of blood. The dentaries' condition suggests that the species may have had an efficient process of immobilizing the fracture during healing, which helped prevent damage to vital blood vessels and nerves. This, along with signs of healing, indicates that the fractures were not imminently fatal.


title: Mosasaurus

text: In 2006, Schulp and colleagues published a study describing a quadrate of "M. hoffmannii" with multiple unnatural openings and an estimated of tissue destroyed. This was likely a severe bone infection initiated by septic arthritis, which progressed to the point where a large portion of the quadrate was reduced to abscess. Extensive amounts of bone reparative tissue were also present, suggesting the infection and subsequent healing process may have progressed for a few months. This level of bone infection would have been tremendously painful and severely hampered the mosasaur's ability to use its jaws. The location of the infection may have also interfered with breathing. Considering how the individual was able to survive such conditions for an extended period of time, Schulp and colleagues speculated it switched to a foraging-type diet of soft-bodied prey like squid that could be swallowed whole to minimize jaw use. The cause of the infection remains unknown, but if it were a result of an intraspecific attack then it is possible one of the openings on the quadrate may have been the point of entry for an attacker's tooth from which the infection entered.


title: Mosasaurus

text: Avascular necrosis has been reported by many studies to be present in every examined specimen of "M. lemonnieri" and "M. conodon". In examinations of "M. conodon" fossils from Alabama and New Jersey and "M. lemonnieri" fossils from Belgium, Rothschild and Martin in 2005 observed that the condition affected between 3-17% of the vertebrae in the mosasaurs' spines. Avascular necrosis is a common result of decompression illness; it involves bone damage caused by the formation of nitrogen bubbles from inhaled air decompressed during frequent deep-diving trips, or by intervals of repetitive diving and short breathing. This indicates that both "Mosasaurus" species may have either been habitual deep-divers or repetitive divers. Agnete Weinreich Carlsen considered it the simplest explanation that such conditions were a product of inadequate anatomical adaptation. Nevertheless, fossils of other mosasaurs with invariable avascular necrosis still exhibit substantial adaptations like eardrums that were well-protected from rapid changes in pressure.


title: Mosasaurus

text: Unnatural fusion of tail vertebrae has been documented in "Mosasaurus", which occurs when the bones remodel themselves after damage from trauma or disease. A 2015 study by Rothschild and Everhart surveyed 15 "Mosasaurus" specimens from North America and Belgium and found cases of fused tail vertebrae in three of them. Two of these cases displayed irregular surface deformities around the fusion site caused by drainage of the vertebral sinuses, which is indicative of a bone infection. The causes of such infections are uncertain, but records of fused vertebrae in other mosasaurs suggest attacks by sharks and other predators as a possible candidate. The third case was determined to be caused by a form of arthritis based on the formation of smooth bridging between fused vertebrae.


title: Mosasaurus

text: It is likely that "Mosasaurus" was viviparous (giving live birth) like most modern mammals today. There is no evidence for live birth in "Mosasaurus" itself, but it is known in a number of other mosasaurs; examples include a skeleton of a pregnant "Carsosaurus", a "Plioplatecarpus" fossil associated with fossils of two mosasaur embryos, and fossils of newborn "Clidastes" from pelagic (open ocean) deposits. Such fossil records, along with a total absence of any evidence suggesting external egg-based reproduction, indicates the likeliness of viviparity in "Mosasaurus". Microanatomical studies on bones of juvenile "Mosasaurus" and related genera have found that their bone structures are comparable to adults. They do not exhibit the bone mass increase found in juvenile primitive mosasauroids to support buoyancy associated with a lifestyle in shallow water, implying that "Mosasaurus" was precocial: they were already efficient swimmers and lived fully functional lifestyles in open water at a very young age, and did not require nursery areas to raise their young. Some areas in Europe and South Dakota have yielded concentrated assemblages of juvenile "M. hoffmannii", "M. missouriensis" and/or "M. lemonnieri". These localities are all shallow ocean deposits, suggesting that juvenile "Mosasaurus" may still have lived in shallow waters.


title: Mosasaurus

text: "Mosasaurus" had a transatlantic distribution, with its fossils having been found in marine deposits on both sides of the Atlantic Ocean. These localities include the Midwest and East Coast of the United States, Canada, Europe, Turkey, Russia, the Levant, the African coastline from Morocco to South Africa, Brazil, Argentina, and Antarctica. During the Late Cretaceous, these regions made up the three seaways inhabited by "Mosasaurus": the Atlantic Ocean, the Western Interior Seaway, and the Mediterranean Tethys. Multiple oceanic climate zones encompassed the seaways, including tropical, subtropical, temperate, and subpolar climates. The wide range of oceanic climates yielded a large diversity of fauna that coexisted with "Mosasaurus".


title: Mosasaurus

text: The Mediterranean Tethys during the Maastrichtian stage was located in what is now Europe, Africa, and the Middle East. In recent studies, the confirmation of paleogeographical affinities extended this range to areas across the Atlantic including Brazil and the East Coast state of New Jersey. It is geographically subdivided into two biogeographic provinces that respectively include the northern and southern Tethyan margins. The two mosasaurs "Mosasaurus" and "Prognathodon" appear to have been the dominant taxa, being widespread and ecologically diversified throughout the seaway.


title: Mosasaurus

text: The northern Tethyan margin was located around the paleolatitudes of 30–40°N, consisting of what is now the European continent, Turkey, and New Jersey. At the time, Europe was a scattering of islands with most of the modern continental landmass being underwater. The margin provided a warm-temperate climate with habitats dominated by mosasaurs and sea turtles. "M. hoffmannii" and "Prognathodon sectorius" were the dominant species in the northern province. In certain areas such as Belgium, other "Mosasaurus" species like "M. lemonnieri" were instead the dominant species, where it's occurrences greatly outnumber those of other large mosasaurs. Other mosasaurs found in the European side of the northern Tethyan margin include smaller genera such as "Halisaurus", "Plioplatecarpus", and "Platecarpus"; the shell-crusher "Carinodens"; and larger mosasaurs of similar trophic levels including "Tylosaurus bernardi" and four other species of "Prognathodon". Sea turtles such as "Allopleurodon hoffmanni" and "Glyptochelone suickerbuycki" were also prevalent in the area and other marine reptiles including indeterminate elasmosaurs have been occasionally found. Marine reptile assemblages in the New Jersey region of the province are generally equivalent with those in Europe; the mosasaur faunae are quite similar but exclude "M. lemonnieri", "Carinodens", "Tylosaurus", and certain species of "Halisaurus" and "Prognathodon". In addition, they exclusively feature "M. conodon", "Halisaurus platyspondylus" and "Prognathodon rapax". Many types of sharks such as "Squalicorax", "Cretalamna", "Serratolamna", and sand sharks, as well as bony fish such as "Cimolichthys", the saber-toothed herring "Enchodus", and the swordfish-like "Protosphyraena" are represented in the northern Tethyan margin.


title: Mosasaurus

text: The southern Tethyan margin was located along the equator between 20°N and 20°S, resulting in warmer tropical climates. Seabeds bordering the cratons in Africa and Arabia and extending to the Levant and Brazil provided vast shallow marine environments. These environments were dominated by mosasaurs and marine side-necked turtles. Of the mosasaurs, "Globidens phosphaticus" is the characteristic species of the southern province; in the African and Arabian domain, "Halisaurus arambourgi" and "Platecarpus ptychodon" were also common mosasaurs alongside "Globidens". "Mosasaurus" was not well-represented: the distribution of "M. beaugei" was restricted to Morocco and Brazil and isolated teeth from Syria suggested a possible presence of "M. lemonnieri", although "M. hoffmannii" also had some presence throughout the province. Other mosasaurs from the southern Tethyan margin include the enigmatic "Goronyosaurus", the shell-crushers "Igdamanosaurus" and "Carinodens", "Eremiasaurus", four other species of "Prognathodon", and various other species of "Halisaurus". Other marine reptiles such as the marine monitor lizard "Pachyvaranus" and the sea snake "Palaeophis" are known there. Aside from "Zarafasaura" in Morocco, plesiosaurs were scarce. As a tropical area, bony fish such as "Enchodus" and "Stratodus" and various sharks were common throughout the southern Tethyan margin.


title: Mosasaurus

text: Many of the earliest fossils of "Mosasaurus" were found in Campanian stage deposits in North America, including the Western Interior Seaway, an inland sea which once flowed through what is now the central United States and Canada, and connected the Arctic Ocean to the modern-day Gulf of Mexico. The region was shallow for a seaway, reaching a maximum depth of about . Extensive drainage from the neighboring continents, Appalachia and Laramidia, brought in vast amounts of sediment. Together with the formation of a nutrient-rich deepwater mass from the mixing of continental freshwater, Arctic waters from the north, and warmer saline Tethyan waters from the south, this created a warm and productive seaway that supported a rich diversity of marine life.


title: Mosasaurus

text: The biogeography of the region has been subdivided into two Interior Subprovinces characterized by different climates and faunal structures, and their borders are separated in modern-day Kansas. The oceanic climate of the Northern Interior Subprovince was likely a cool temperate one, while the Southern Interior Subprovince had warm temperate to subtropical climates. The fossil assemblages throughout these regions suggest a complete faunal turnover when "M. missouriensis" and "M. conodon" appeared at 79.5 Ma, indicating that the presence of "Mosasaurus" in the Western Interior Seaway had a profound impact on the restructuring of marine ecosystems. The faunal structure of both provinces was generally much more diverse prior to the appearance of "Mosasaurus", during a faunal stage known as the Niobraran Age, than it was during the following Navesinkan Age.


title: Mosasaurus

text: In what is now Alabama within the Southern Interior Subprovince, most of the key genera including sharks like "Cretoxyrhina" and the mosasaurs "Clidastes", "Tylosaurus", "Globidens", "Halisaurus", and "Platecarpus" disappeared and were replaced by "Mosasaurus". During the Navesinkan Age, "Mosasaurus" dominated the whole region, accounting for around two-thirds of all mosasaur diversity with "Plioplatecarpus" and "Prognathodon" sharing the remaining third. The Northern Interior Subprovince also saw a restructuring of mosasaur assemblages, characterized by the disappearance of mosasaurs like "Platecarpus" and their replacement by "Mosasaurus" and "Plioplatecarpus". Some Niobraran genera such as "Tylosaurus", "Cretoxyrhina", hesperornithids, and plesiosaurs including elasmosaurs such as "Terminonatator" and polycotylids like "Dolichorhynchops" maintained their presence until around the end of the Campanian, during which the entire Western Interior Seaway started receding from the north. "Mosasaurus" continued to be the dominant genus in the seaway until the end of the Navesinkan Age at the end of the Cretaceous. Contemporaneous fauna included sea turtles such as "Protostega" and "Archelon"; many species of sea birds including "Baptornis", "Ichthyornis", and "Halimornis"; sharks such as the mackerel sharks "Cretalamna", "Squalicorax", "Pseudocorax", and "Serratolamna", the goblin shark "Scapanorhynchus", the sand tiger "Odontaspis", and the sawfish-like "Ischyrhiza"; and bony fish such as "Enchodus", "Protosphyraena", "Stratodus", and the ichthyodectids "Xiphactinus" and "Saurodon".


title: Mosasaurus

text: "Mosasaurus" is known from late Maastrichtian deposits in the Antarctic Peninsula, specifically the López de Bertodano Formation in Seymour Island. Located within the polar circle at around 65°S, temperatures at medium to large water depths would have been around on average, while sea surface temperatures may have dropped below freezing and sea ice may have formed at times. "Mosasaurus" appears to be the most diverse mosasaur in the Maastrichtian Antarctica. At least two species of "Mosasaurus" have been described, but the true number of species is unknown as remains are often fragmentary and specimens are described in open nomenclature. These species include one comparable with "M. lemonnieri", and another that appears to be closely related to "M. hoffmannii". "M. sp." has also been described. However, it is possible that such specimens may actually represent "Moanasaurus", although this depends on the outcome of a pending revision of the genus. At least four other mosasaur genera have been reported in Antarctica, including "Plioplatecarpus", the mosasaurines "Moanasaurus" and "Liodon", and "Kaikaifilu". The validity of some of these genera is disputed as they are primarily based on isolated teeth. "Prognathodon" and "Globidens" are also expected to be present based on distribution trends of both genera, although conclusive fossils have yet to be found. Other Antarctic marine reptiles included elasmosaurid plesiosaurs like "Aristonectes" and another indeterminate elasmosaurid. The fish assemblage of the López de Bertodano Formation was dominated by "Enchodus" and ichthyodectiformes.


title: Mosasaurus

text: Known fossils of "Mosasaurus" have typically been recovered from deposits representing nearshore habitats during the Cretaceous period, with some fossils coming from deeper-water deposits. Lingham-Soliar (1995) elaborated on this, finding that Maastrichtian deposits in the Netherlands with "M. hoffmannii" occurrences represented nearshore waters around deep. Changing temperatures and an abundance in marine life were characteristic of these localities. The morphological build of "M. hoffmannii", nevertheless, was best adapted for a pelagic surface lifestyle.