title: Jude Bellingham

text: Jude Victor William Bellingham (born 29 June 2003) is an English professional footballer who plays as a midfielder for Bundesliga club Borussia Dortmund and the England national team.


title: Jude Bellingham

text: Bellingham joined Birmingham City as an under-8, became the club's youngest ever first-team player when he made his senior debut in August 2019, at the age of , and played regularly during the 2019–20 season. He joined Borussia Dortmund in July 2020, and in his first competitive match became their youngest ever goalscorer.


title: Jude Bellingham

text: He represented England at under-15, under-16, under-17 and under-21 levels. He made his first appearance for the senior team in November 2020, and represented the country at UEFA Euro 2020.


title: Jude Bellingham

text: Jude Victor William Bellingham was born on 29 June 2003 in Stourbridge, in the Metropolitan Borough of Dudley, West Midlands, the eldest son of Denise and Mark Bellingham. He is of Irish descent through a paternal grandparent. His father Mark is a sergeant in the West Midlands Police and was a prolific goalscorer in non-League football. Bellingham's younger brother, Jobe, is a Birmingham City player. Bellingham attended Priory School in Edgbaston, Birmingham.


title: Jude Bellingham

text: Bellingham joined Birmingham City as an under-8, after playing for Stourbridge. He played for their under-18 team at 14, and made his debut for their under-23 team at the age of 15, on 15 October 2018 away to Nottingham Forest's U23s. Entering the game after an hour, he scored the only goal in the 87th minute "sliding in to force the ball over the goal line after pressure by Kyle McFarlane on the keeper diverted the ball into his path." By March 2019, he had three goals from ten development squad appearances, had featured in "FourFourTwo" list of the "50 most exciting teenagers in English football", and was mentioned as of interest to major European clubs. He was gradually introduced to the first-team environment while still a schoolboy: increasingly training with the seniors, he accompanied them on matchday to observe, and travelled as the "19th man" for a Championship match in March.


title: Jude Bellingham

text: Bellingham took up a two-year scholarship with Birmingham City to begin in July 2019. He was part of the first-team training camp in Portugal, played and scored in pre-season friendlies, and was given squad number 22 for the 2019–20 season. On 6 August, when he started the EFL Cup first round visit to Portsmouth, Bellingham became Birmingham City's youngest ever first-team player. At , he lowered the record set by Trevor Francis in 1970 by 101 days. He played for 80 minutes in the 3–0 defeat, and was the "Birmingham Mail" man of the match. He made his first Football League appearance 19 days later, as a second-half substitute in a 3–0 defeat away to Swansea City, and his home debut on 31 August against Stoke City. Replacing the injured Jefferson Montero after half an hour, Bellingham scored the winneralbeit via a generous deflectionas Birmingham came back from 1–0 down to beat Stoke 2–1, and thus became their youngest ever goalscorer, aged 16 years and 63 days. He started the next match, away to Charlton Athletic two weeks later, and scored the only goal from Kerim Mrabti's cutback.


title: Jude Bellingham

text: Bellingham continued as a permanent fixture in the matchday squad, sometimes as a substitute but mainly in the starting eleven. He was eased into the team on the left wing, moved into central midfield "where he could gain more confidence", and then used "in a more advanced role" once the staff were sure he could cope with the responsibility. He was EFL Young Player of the Month for November 2019. According to head coach Pep Clotet, Bellingham himself "feels more comfortable in midfield, and more comfortable when he can get closer to the opposition box."


title: Jude Bellingham

text: He was linked with moves to numerous major clubs in January 2020; on deadline day, Birmingham were reported to have turned down a £20 million bid from Manchester United. Bellingham continued as a first-team regular, and by the time the season was suspended because of the COVID-19 pandemic, he had made 32 league appearances. He remained an integral part of the team once the season resumed behind closed doors, and set up a late equaliser for Lukas Jutkiewicz against Charlton Athletic that made Birmingham's league position less precarious with two matches still to play. He finished the season with four goals from 44 appearances in all competitions, 41 in the league, as Birmingham avoided relegation despite losing the last match of the season. In appreciation of what Bellingham achieved in such a short time with the first team, the club announced that they would retire his number 22 shirt, "to remember one of our own and to inspire others." At the EFL Awards, he was named both Championship Apprentice of the Year and EFL Young Player of the Season.


title: Jude Bellingham

text: It had long become obvious that Bellingham would leave Birmingham, and it was reported that he and his father had visited several major clubs, of which Manchester United and Bundesliga club Borussia Dortmund were the favourites. Impressed by Dortmund's record of including young players as regulars in the first team, as evidenced by the likes of Jadon Sancho, that became his destination of choice. He flew to Germany for a medical, and the transfer was confirmed on 20 July 2020: he would join after Birmingham's last match of the season. The undisclosed fee was understood by Sky Sports to be an initial £25 millionmaking him the most expensive 17-year-old in historyplus "several million more" dependent on performance-related criteria.


title: Jude Bellingham

text: Bellingham made his debut on 14 September 2020, starting the first match of Dortmund's 2020–21 season against third-tier MSV Duisburg in the DFB-Pokal, aged . After half an hour, he scored the second goal in a 5–0 win, becoming the club's youngest goalscorer in the DFB-Pokal, taking six days off Giovanni Reyna's record, as well as their youngest scorer in any competitive match, breaking Nuri Şahin's record by five days. Five days later, he marked his league debut with the assist for Reyna's opening goal in a 3–0 win over Borussia Mönchengladbach, and was named as Bundesliga Rookie of the Month for September. When Bellingham faced Lazio in the group stage on 20 October, aged 17 years and 113 days, he became the youngest Englishman to start a Champions League match, breaking the record previously set by Phil Foden.


title: Jude Bellingham

text: In the first three months of the season, Bellingham was a regular in all competitions, with six starts and seven substitute appearances in the Bundesliga as well as four Champions League starts. He missed the first two matches of 2021 with a foot injury, but returned to action as an increasingly regular starter. He was involved in Marco Reus's goal in the away leg of the Champions League quarter-final against Manchester City He scored against Man City early in the second leg, but Dortmund could not retain their away-goal advantage. In between, Bellingham scored his first Bundesliga goal from Reyna's knock-down to equalise with VfB Stuttgart early in the second half; Dortmund won 3–2. Bellingham started for Dortmund in their 4–1 victory over RB Leipzig in the 2021 DFB-Pokal Final. He was booked in the first half and replaced by Thorgan Hazard at half-time with his team 3–0 ahead. He finished the season with 29 appearances and one goal in the Bundesliga, 46 appearances and four goals in all competitions, and was voted Newcomer of the Season by his fellow players.


title: Jude Bellingham

text: Bellingham was runner-up to Pedri of Barcelona in the 2021 Kopa Trophy, awarded to the best under-21 male player as voted by previous winners of the Ballon d'Or. On 4 December, Bellingham played in "Der Klassiker" against Bayern Munich. He made the assists for both Dortmund goals, but Bayern won the match 3–2 via a 77th-minute penalty awarded after lengthy VAR involvement. Earlier in the fixture, two Dortmund penalty appeals were turned down by referee Felix Zwayer, who refused to review either. Interviewed live by Viaplay immediately after the match, Bellingham was critical of Zwayer's decisions, and made reference to his part in the 2005 German football match-fixing scandal, saying: "You give a referee, that has match fixed before, the biggest game in Germany. What do you expect?" The DFB wrote to Bellingham asking for his comments as a matter of urgency. He was later fined €40,000 by the German Football Association.


title: Jude Bellingham

text: Bellingham made his England under-15 debut against Turkey in December 2016. In recognition of his captaining that team during the 2017–18 season, he was presented with a Special Achievement Award at the 2018 Birmingham City Academy awards night. By the end of 2018 he had made his first appearance for the England under-16 team, and went on to feature in eleven games, score four goals, and captain the team. He was included in England's under-17 squad for the Syrenka Cup, a friendly tournament held in September 2019 in preparation for the 2020 European Championship qualifiers the following March. He made his debut as a substitute in England's opening match of the tournament, a 5–0 win over Finland in which he scored the third goal, and captained the team in their second fixture, in which they came back from a goal behind to beat Austria 4–2 and qualify for the final. Again, Bellingham scored the third goal. He retained the captaincy for the final, in which England beat hosts Poland on penalties following a 2–2 draw, and was named player of the tournament.


title: Jude Bellingham

text: Bellingham received his first call-up to the under-21 squad for European Championship qualifiers against Kosovo and Austria in September 2020. He became the youngest player to appear for England U21 when he came on to replace Tom Davies after 62 minutes of the match against Kosovo on 4 September with England 3–0 ahead, and scored after 85 minutes to complete the 6–0 victory.


title: Jude Bellingham

text: In November 2020, after James Ward-Prowse and Trent Alexander-Arnold withdrew through injury, Bellingham was called up to the England senior squad for the first time. He made his debut in a friendly against the Republic of Ireland at Wembley on 12 November, replacing Mason Mount after 73 minutes of a 3–0 win. At , he became England's third-youngest full international; only Theo Walcott and Wayne Rooney had appeared at a younger age. Bellingham was named in the England squad for the UEFA Euro 2020 tournament, which was delayed until June 2021 because of the COVID-19 pandemic. When he came on as an 82nd-minute substitute in England's opening match, a 1–0 win over Croatia at Wembley on 13 June, aged 17 years and 349 days, he became both the youngest Englishman to play at any major tournament and the youngest of any nationality to play at a European Championship; the latter record was broken by Poland's Kacper Kozłowski just six days later.


title: Jude Bellingham

text: Bellingham's first senior international goal, a header from Luke Shaw's cross to open the scoring in England's 6–2 win over Iran in their first game of the 2022 World Cup on 21 November 2022, made him the second youngest scorer for England at a World Cup. He also made the run and pass to Harry Kane who crossed for Raheem Sterling to score England's third goal, and played the through ball from which Callum Wilson set up their sixth for Jack Grealish. He then followed this up during the game against Senegal with a run through Senegal's defence, assisting Jordan Henderson's goal in the 39th minute. He then played a key role in Harry Kane's goal in the 3rd minute of extra time in the first half, setting up Phil Foden to make the assist.


title: Rhea Perlman

text: Rhea Jo Perlman (born March 31, 1948) is an American actress. She played head-waitress Carla Tortelli in the sitcom "Cheers" (1982–1993). Over the course of 11 seasons, Perlman was nominated for ten Emmy Awards for Outstanding Supporting Actress – winning four times – and was nominated for a record seven Golden Globe Awards for Best Supporting Actress in a Television Series. She has also appeared in films, including "Canadian Bacon" (1995), "Matilda" (1996), "The Sessions" (2012), and "Poms" (2019).


title: Rhea Perlman

text: Perlman was born on March 31, 1948, in Coney Island, Brooklyn, to Philip Perlman, a Polish immigrant who was a manager at a doll parts factory and Adele, a bookkeeper. She grew up in Bensonhurst in a Jewish family with additional roots in Russia. She has a sister, Heide, who is a television writer, story editor, and producer who worked on "Cheers", "Frasier" and "The Tracey Ullman Show". In the mid-1980s, her parents moved to Los Angeles, and her father became an extra on "Cheers". His character became known by his real name, Phil, and he managed to get a few lines over the years as he appeared in more than 30 episodes. He created a second career as a character actor, appearing in several films and television shows, including "Throw Momma from the Train", "Hoffa", and "Frasier".


title: Rhea Perlman

text: Perlman began her acting career with a small role as an attendant in the off-off-Broadway play "Dracula Sabbat", which ran from September 1970 to June 1971. In 1972, she played a bit role in the film "Hot Dogs for Gauguin". That same year she appeared in Westbeth Playwrights Feminist Collective's production of "Up – An Uppity Revue", along with her future husband, Danny DeVito.


title: Rhea Perlman

text: One of her first notable parts was a recurring role on the television show "Taxi" as Zena, the sweet girlfriend of Louie De Palma (played by DeVito).


title: Rhea Perlman

text: Following that, she had a role in a small play portraying a much tougher character. Producers Glen and Les Charles saw her in that play, which led to her landing the role as wisecracking barmaid Carla Tortelli on their sitcom "Cheers" in 1982. The series struggled with ratings in its first season, but by the time it ended in 1993, it was one of the most popular and successful shows of all time, winning 20 Emmy awards out of 95 nominations.


title: Rhea Perlman

text: Perlman won the Emmy Award for Outstanding Supporting Actress in a Comedy four times: in 1984, 1985, 1986 and 1989. Over her 11 seasons on "Cheers", she was nominated for an Emmy every year but 1992, becoming the "Cheers" star to have the most wins and nominations. She was also nominated for a Golden Globe for Best Supporting Actress six times, more than anyone else in that category. In 2011, CBS named Carla Tortelli as one of the greatest TV characters of all time.


title: Rhea Perlman

text: In 1986, Perlman starred in an episode of Steven Spielberg's "Amazing Stories" titled "The Wedding Ring," which also starred DeVito as her character's husband.


title: Rhea Perlman

text: In the 1990s, Perlman starred in several TV movies and motion pictures. In 1992, she starred in the made-for-TV-movie, "To Grandmother's House We Go" opposite Ashley Olsen and Mary-Kate Olsen, playing the wife of Jerry Van Dyke's character; the couple kidnapped the Olsen Twins' characters, hoping to cash in on ransom before Christmas. Other TV films in which she starred included the dramas "A Place to Be Loved" and "In Spite of Love."


title: Rhea Perlman

text: Perlman's motion picture roles included "There Goes The Neighborhood" (1992), "Canadian Bacon" (1995), "Carpool" (1996), "Sunset Park" (1996), and "Matilda" (1996). She had a cameo in the film "10 Items Or Less" (2006), and also starred in the 2007 independent film "Love Comes Lately."


title: Rhea Perlman

text: In 1994, Perlman voiced 9-Eye in The Timekeeper, a Circle-Vision show at the Magic Kingdom in Tomorrowland. She later starred in the 1996 sitcom "Pearl" as the title character and was featured on the 2001 TV drama "Kate Brasher." Among her notable guest appearances was on the fourth-season premiere of "Becker", which starred "Cheers" co-star Ted Danson.


title: Rhea Perlman

text: Perlman also appeared in a 2000 television film "," in which she impersonated Jacqueline Kennedy. She also portrayed a therapist called Dr. Parella in the 2000 film "Secret Cutting," which follows the story of a young girl named Dawn who self-injures. In 2007, Perlman appeared as Bertha in the West End of London in the comedy "Boeing Boeing." In 2008, she starred in the Hallmark Channel original movie, "The Christmas Choir," and appeared in "Beethoven's Big Break" in 2008 as Patricia Benji. In 2009, she appeared as Tanya's mother on the series "Hung" for Home Box Office Networks. In 2011, Perlman had a guest appearance as Mittens in one episode of "Wilfred."


title: Rhea Perlman

text: In 2009, Perlman and her daughter Lucy DeVito starred in the off-Broadway play "Love, Loss, and What I Wore", adapted by Nora and Delia Ephron, at the Westside Theatre.


title: Rhea Perlman

text: From 2014 to 2017, she starred in a recurring role on "The Mindy Project" as Danny's mother, Annette Castellano.


title: Rhea Perlman

text: Perlman is the author of the illustrated children's book series "Otto Undercover," whose six books to date (as of the middle of May 2012) are "Born to Drive," "Canyon Catastrophe," "Water Balloon Doom," "Toxic Taffy Takeover," "The Brink of Ex-stink-tion," and "Brain Freeze."


title: Rhea Perlman

text: Perlman met Danny DeVito on January 17, 1971, when she went to see a friend in the single performance of the play "The Shrinking Bride", which also featured DeVito. They moved in together two weeks after meeting and married on January 28, 1982. They have three children: Lucy Chet DeVito, Grace Fan DeVito, and Jacob Daniel DeVito. Perlman, who is Jewish, and DeVito, who was raised Catholic, raised their children celebrating the major holidays of both religions but did not give their children any religious identity. Perlman told the "Los Angeles Times" in 1998: "We do all the holidays to keep the traditions and the culture going, but I truly don't have a great feeling about any particular organized religion, and I don't think it's right to impose one on my kids. I feel like I'm bringing them up to be good people, and that's what it's about."


title: Rhea Perlman

text: The family resided in Beverly Hills, California, and they owned a vacation home in Interlaken, New Jersey. Throughout their relationship, Perlman and DeVito have acted alongside each other several times, including in the TV show "Taxi" and the feature film "Matilda".


title: Rhea Perlman

text: Perlman and DeVito separated in October 2012. However, in March 2013, it was reported that they had reconciled. The couple later separated again for a second time in March 2017 on amicable terms. Although the two no longer live together, Perlman said she has no intention of divorcing DeVito. In 2019, Perlman told interviewer Andy Cohen that she and DeVito have become closer friends after their separation than they were in their final years as a couple.


title: Black Dahlia

text: Elizabeth Short (July 29, 1924 – January 14–15, 1947), known posthumously as the Black Dahlia, was an American woman found murdered in the Leimert Park neighborhood of Los Angeles on January 15, 1947. Her case became highly publicized owing to the gruesome nature of the crime, which included the mutilation of her corpse, which was bisected at the waist.


title: Black Dahlia

text: A native of Boston, Short spent her early life in New England and Florida before relocating to California, where her father lived. It is commonly held that Short was an aspiring actress, though she had no known acting credits or jobs during her time in Los Angeles. She would acquire the nickname of the Black Dahlia posthumously, as newspapers of the period often nicknamed particularly lurid crimes; the term may have originated from a film noir murder mystery, "The Blue Dahlia", released in 1946. After the discovery of her body, the Los Angeles Police Department (LAPD) began an extensive investigation that produced over 150 suspects but yielded no arrests.


title: Black Dahlia

text: Short's unsolved murder and the details surrounding it have had a lasting cultural intrigue, generating various theories and public speculation. Her life and death have been the basis of numerous books and films, and her murder is frequently cited as one of the most famous unsolved murders in American history, as well as one of the oldest unsolved cases in Los Angeles County. It has likewise been credited by historians as one of the first major crimes in post–World War II America to capture national attention.


title: Black Dahlia

text: Elizabeth Short was born on July 29, 1924, in the Hyde Park section of Boston, Massachusetts, the third of five daughters of Cleo A. Short and wife Phoebe May Sawyer. In 1927 the Short family briefly relocated to Portland, Maine, before settling in Medford, a suburb of Boston, that same year. Short's father built miniature golf courses until he lost most of his savings in the 1929 stock market crash. In 1930, his car was found abandoned on the Charlestown Bridge, and it was assumed that he had jumped into the Charles River. Believing her husband to be deceased, Short's mother began working as a bookkeeper to support the family.


title: Black Dahlia

text: Troubled by bronchitis and severe asthma attacks, Short underwent lung surgery at age 15, after which doctors suggested she periodically relocate to a milder climate to prevent further respiratory problems. Short's mother sent her to spend winters in Miami, Florida, with family friends for the next three years. In her sophomore year, Short dropped out of Medford High School.


title: Black Dahlia

text: In late 1942, Short's mother received a letter of apology from her presumed-deceased husband, which revealed that he was in fact alive and had started a new life in California. In December, at age 18, Short relocated to Vallejo, California, to live with her father, whom she had not seen since age 6. At the time he was working at the nearby Mare Island Naval Shipyard on San Francisco Bay. Arguments between Short and her father led to her moving out in January 1943.


title: Black Dahlia

text: Short took a job at the Base Exchange at Camp Cooke (now Vandenberg Air Force Base) near Lompoc, briefly living with a U.S. Army Air Force sergeant who reportedly abused her. She left Lompoc in mid-1943 and moved to Santa Barbara, where she was arrested on September 23, 1943, for drinking at a local bar while underage. The juvenile authorities sent her back to Massachusetts but she returned instead to Florida, making only occasional visits to her family near Boston.


title: Black Dahlia

text: While in Florida, Short met Major Matthew Michael Gordon, Jr., a decorated Army Air Force officer of the 2nd Air Commando Group, who was training for deployment to Southeast Asian theater of World War II. Short later told friends that Gordon had written to propose marriage while he was recovering from injuries from a plane crash in India. She accepted his offer, but Gordon died in a second crash on August 10, 1945, less than a week before the end of the war.


title: Black Dahlia

text: In July 1946, Short relocated to Los Angeles to visit Army Air Force Lieutenant Joseph Gordon Fickling, an acquaintance from Florida, who was stationed at the Naval Reserve Air Base in Long Beach. Short spent the last six months of her life in southern California, mostly in the Los Angeles area; shortly before her death she had been working as a waitress, and rented a room behind the Florentine Gardens nightclub on Hollywood Boulevard. She has been variously described and depicted as an aspiring or "would-be" actress. According to some sources, she did in fact have aspirations to be a film star, though she had no known acting jobs or credits.


title: Black Dahlia

text: On January 9, 1947, Short returned to her home in Los Angeles after a brief trip to San Diego with Robert "Red" Manley, a 25-year-old married salesman she had been dating. Manley stated that he dropped Short off at the Biltmore Hotel in downtown Los Angeles, and that Short was to meet her sister, who was visiting from Boston, that afternoon. By some accounts, staff of the Biltmore recalled having seen Short using the lobby telephone. Shortly after, she was allegedly seen by patrons of the Crown Grill Cocktail Lounge at 754 South Olive Street, approximately away from the Biltmore.


title: Black Dahlia

text: On the morning of January 15, 1947, Short's naked body, severed into two pieces, was found on a vacant lot on the west side of South Norton Avenue, midway between Coliseum Street and West 39th Street (at ) in the neighborhood of Leimert Park. At the time, Leimert Park was largely undeveloped. Local resident Betty Bersinger discovered the body at approximately 10 a.m. while walking with her three-year-old daughter, initially thinking she had found a discarded store mannequin. When she realized it was a corpse, she rushed to a nearby house and telephoned the police.


title: Black Dahlia

text: Short's severely mutilated body was completely severed at the waist and drained of blood, leaving her skin a pallid white. Medical examiners determined that she had been dead for around ten hours prior to the discovery, leaving her time of death either sometime during the evening of January 14 or the early morning hours of January 15. The body had apparently been washed by the killer. Short's face had been slashed from the corners of her mouth to her ears, creating an effect known as the "Glasgow smile". She had several cuts on her thigh and breasts, where entire portions of flesh had been sliced away. The lower half of her body was positioned a foot away from the upper, and her intestines had been tucked neatly beneath her buttocks. The corpse had been "posed", with her hands over her head, her elbows bent at right angles, and her legs spread apart.


title: Black Dahlia

text: Upon the discovery, a crowd of passersby and reporters began to gather; "Los Angeles Herald-Express" reporter Aggie Underwood was among the first to arrive at the scene, and took several photos of the corpse and crime scene. Near the body, detectives located a heel print on the ground amid the tire tracks, and a cement sack containing watery blood was also found nearby.


title: Black Dahlia

text: An autopsy of Short's body was performed on January 16, 1947, by Frederick Newbarr, the Los Angeles County coroner. Newbarr's autopsy report stated that Short was tall, weighed , and had light blue eyes, brown hair, and badly decayed teeth. There were ligature marks on her ankles, wrists, and neck, and an "irregular laceration with superficial tissue loss" on her right breast. Newbarr also noted superficial lacerations on the right forearm, left upper arm, and the lower left side of the chest.


title: Black Dahlia

text: The body had been cut completely in half by a technique taught in the 1930s called a hemicorporectomy. The lower half of her body had been removed by transecting the lumbar spine between the second and third lumbar vertebrae, thus severing the intestine at the duodenum. Newbarr's report noted "very little" ecchymosis (bruising) along the incision line, suggesting it had been performed after death. Another "gaping laceration" measuring in length ran longitudinally from the umbilicus to the suprapubic region. The lacerations on each side of the face, which extended from the corners of the lips, were measured at on the right side of the face, and on the left. The skull was not fractured, but there was bruising noted on the front and right side of her scalp, with a small amount of bleeding in the subarachnoid space on the right side, consistent with blows to the head. The cause of death was determined to be hemorrhaging from the lacerations to her face and the shock from blows to the head and face. Newbarr noted that Short's anal canal was dilated at , suggesting that she might have been raped. Samples were taken from her body testing for the presence of sperm, but the results came back negative.


title: Black Dahlia

text: Short was identified after her fingerprints were sent to the FBI via Soundphoto, a device which transmitted images by telephone and was normally used for news photographs; Short's fingerprints were on file from her 1943 arrest. Immediately following Short's identification, reporters from William Randolph Hearst's "Los Angeles Examiner" contacted her mother, Phoebe Short, in Boston, and told her that her daughter had won a beauty contest. It was only after prying as much personal information as they could from Phoebe that the reporters revealed that her daughter had in fact been murdered. The newspaper offered to pay her airfare and accommodations if she would travel to Los Angeles to help with the police investigation. That was yet another ploy since the newspaper kept her away from police and other reporters to protect its scoop. The "Examiner" and another Hearst newspaper, the "Los Angeles Herald-Express", later sensationalized the case, with one article from the "Examiner" describing the black tailored suit Short was last seen wearing as "a tight skirt and a sheer blouse". The media nicknamed her the "Black Dahlia", and described her as an "adventuress" who "prowled Hollywood Boulevard". Additional newspaper reports, such as one published in the "Los Angeles Times" on January 17, deemed the murder a "sex fiend slaying".


title: Black Dahlia

text: On January 21, 1947, a person claiming to be Short's killer placed a phone call to the office of James Richardson, the editor of the "Examiner", congratulating Richardson on the newspaper's coverage of the case, and stated he planned on eventually turning himself in, but not before allowing police to pursue him further. Additionally, the caller told Richardson to "expect some souvenirs of Beth Short in the mail".


title: Black Dahlia

text: On January 24, a suspicious manila envelope was discovered by a U.S. Postal Service worker. The envelope had been addressed to "The "Los Angeles Examiner" and other Los Angeles papers" with individual words that had been cut-and-pasted from newspaper clippings; additionally, a large message on the face of the envelope read: "Here is Dahlia's belongings [,] letter to follow". The envelope contained Short's birth certificate, business cards, photographs, names written on pieces of paper, and an address book with the name Mark Hansen embossed on the cover. The packet had been carefully cleaned with gasoline, similarly to Short's body, which led police to suspect the packet had been sent directly by her killer. Despite the efforts to clean the packet, several partial fingerprints were lifted from the envelope and sent to the Federal Bureau of Investigation for testing; however, the prints were compromised in transit and thus could not be properly analyzed. The same day the packet was received by the "Examiner", a handbag and a black suede shoe were reported to have been seen on top of a garbage can in an alley a short distance from Norton Avenue, from where Short's body had been discovered. The items were recovered by police, but they had also been wiped clean with gasoline, destroying any fingerprints.


title: Black Dahlia

text: On March 14, an apparent suicide note scrawled in pencil on a bit of paper was found tucked in a shoe in a pile of men's clothing by the ocean's edge at the foot of Breeze Ave., Venice. The note read: "To whom it may concern: I have waited for the police to capture me for the Black Dahlia killing, but have not. I am too much of a coward to turn myself in, so this is the best way out for me. I couldn't help myself for that, or this. Sorry, Mary." The pile of clothing was first seen by a beach caretaker, who reported the discovery to John Dillon, lifeguard captain. Dillon immediately notified Capt. L. E. Christensen of West Los Angeles Police Station. The clothes included a coat and trousers of blue herringbone tweed, a brown and white T shirt, white jockey shorts, tan socks and tan moccasin leisure shoes, size about eight. The clothes gave no clue about the identity of their owner.


title: Black Dahlia

text: Police quickly deemed Mark Hansen, the owner of the address book found in the packet, a suspect. Hansen was a wealthy local nightclub and theater owner and an acquaintance at whose home Short had stayed with friends, and according to some sources, he also confirmed that the purse and shoe discovered in the alley were in fact Short's. Ann Toth, Short's friend and roommate, told investigators that Short had recently rejected sexual advances from Hansen, and suggested it as potential cause for him to kill her; however, he was cleared of suspicion in the case. In addition to Hansen, the Los Angeles Police Department interviewed over 150 men in the ensuing weeks whom they believed to be potential suspects. Manley, who had been one of the last people to see Short alive, was also investigated, but was cleared of suspicion after passing numerous polygraph examinations. Police also interviewed several persons found listed in Hansen's address book, including Martin Lewis, who had been an acquaintance of Short's. Lewis was able to provide an alibi for the date of Short's murder, as he was in Portland, Oregon, visiting his father-in-law, who was dying of kidney failure.


title: Black Dahlia

text: A total of 750 investigators from the LAPD and other departments worked on the case during its initial stages, including 400 sheriff's deputies and 250 California State Patrol officers. Various locations were searched for potential evidence, including storm drains throughout Los Angeles, abandoned structures, and various sites along the Los Angeles River, but the searches yielded no further evidence. City councilman Lloyd G. Davis posted a reward for information leading police to Short's killer. After the announcement of the reward, various persons came forward with confessions, most of which police dismissed as false. Several of the false confessors were charged with obstruction of justice.


title: Black Dahlia

text: On January 26, another letter was received by the "Examiner", this time handwritten, which read: "Here it is. Turning in Wed., Jan. 29, 10 am. Had my fun at police. Black Dahlia Avenger". The letter also named a location at which the supposed killer would turn himself in. Police waited at the location on the morning of January 29, but the alleged killer did not appear. Instead, at 1:00pm, the "Examiner" offices received another cut-and-pasted letter, which read: "Have changed my mind. You would not give me a square deal. Dahlia killing was justified."


title: Black Dahlia

text: The graphic nature of the crime and the subsequent letters received by the "Examiner" had resulted in a media frenzy surrounding Short's murder. Both local and national publications covered the story heavily, many of which reprinted sensationalistic reports suggesting that Short had been tortured for hours prior to her death; the information, however, was false, yet police allowed the reports to circulate so as to conceal Short's true cause of death—cerebral hemorrhage—from the public. Further reports about Short's personal life were publicized, including details about her alleged declining of Hansen's romantic advances; additionally, a stripper who was an acquaintance of Short's told police that she "liked to get guys worked up over her, but she'd leave them hanging dry." This led some reporters (namely the "Herald-Express"s Bevo Means) and detectives to look into the possibility that Short was a lesbian, and begin questioning employees and patrons of gay bars in Los Angeles; this claim, however, remained unsubstantiated. The "Herald-Express" also received several letters from the purported killer, again made with cut-and-pasted clippings, one of which read: "I will give up on Dahlia killing if I get 10 years. Don't try to find me."


title: Black Dahlia

text: On February 1, the "Los Angeles Daily News" reported that the case had "run into a Stone Wall", with no new leads for investigators to pursue. The "Examiner" continued to run stories on the murder and the investigation, which was front-page news for 35 days following the discovery of the body.


title: Black Dahlia

text: When interviewed, lead investigator Captain Jack Donahue told the press that he believed Short's murder had taken place in a remote building or shack on the outskirts of Los Angeles, and her body transported into the city where it was disposed of. Based on the precise cuts and dissection of Short's corpse, the LAPD looked into the possibility that the murderer had been a surgeon, doctor, or someone with medical knowledge. In mid-February 1947, the LAPD served a warrant to the University of Southern California Medical School, which was located near the site where Short's body had been discovered, requesting a complete list of the program's students. The university agreed so long as the students' identities remained private. Background checks were conducted, but yielded no results.


title: Black Dahlia

text: By the spring of 1947, Short's murder had become a cold case with few new leads. Sergeant Finis Brown, one of the lead detectives on the case, blamed the press for compromising the investigation through reporters' probing of details and unverified reporting. In September 1949, a grand jury convened to discuss inadequacies in the LAPD's homicide unit based on their failure to solve numerous murders—especially those of women and children—in the past several years, Short's being one of them. In the aftermath of the grand jury, further investigation was done on Short's past, with detectives tracing her movements between Massachusetts, California, and Florida, and also interviewed people who knew her in Texas and New Orleans. However, the interviews yielded no useful information in the murder.


title: Black Dahlia

text: The notoriety of Short's murder has spurred a large number of confessions over the years, many of which have been deemed false. During the initial investigation into her murder, police received a total of 60 confessions, most made by men. Since that time, over 500 people have confessed to the crime, some of whom had not even been born at the time of her death. Sergeant John P. St. John, a detective who worked the case until his retirement, stated, "It is amazing how many people offer up a relative as the killer."


title: Black Dahlia

text: In 2003, Ralph Asdel, one of the original detectives on the case, told the "Times" that he believed he had interviewed Short's killer, a man who had been seen with his sedan parked near the vacant lot where her body was discovered in the early morning hours of January 15, 1947. A neighbor driving by that day stopped to dispose of a bag of lawn clippings in the vacant lot when he saw a parked sedan, allegedly with its right rear door open; the driver of the sedan was standing in the lot. His arrival apparently startled the owner of the sedan, who approached his car and peered in the window before returning to the sedan and driving away. The owner of the sedan was followed to a local restaurant where he worked, but was ultimately cleared of suspicion.


title: Black Dahlia

text: Suspects remaining under discussion by various authors and experts include a doctor named Walter Bayley, proposed by the former "Times" copyeditor Larry Harnisch; "Times" publisher Norman Chandler, whom biographer Donald Wolfe claims impregnated Short; Leslie Dillon, Joseph A. Dumais, Artie Lane (a.k.a. Jeff Connors), Mark Hansen, Dr. Francis E. Sweeney, Woody Guthrie, Bugsy Siegel, Orson Welles, George Hodel, Hodel's friend Fred Sexton, George Knowlton, Robert M. "Red" Manley, Patrick S. O'Reilly, and Jack Anderson Wilson.


title: Black Dahlia

text: George Hill Hodel Jr. was a suspect; like the others, he was never formally charged with the crime. He came to wider attention as a suspect after his death when he was accused by his son, Los Angeles homicide detective Steve Hodel, of killing Short and committing several additional murders. Prior to the Dahlia case, he was also a suspect in the death of his secretary, Ruth Spaulding, but was not charged; and was accused of raping his own daughter, Tamar, but acquitted. He fled the country several times, and spent 1950 to 1990 in the Philippines.


title: Black Dahlia

text: Several crime authors, as well as Cleveland detective Peter Merylo, have suspected a link between the Short murder and the Cleveland Torso Murders, which took place in Cleveland, Ohio, between 1934 and 1938. As part of their investigation into other murders that took place before and after the Short killing, the original LAPD investigators studied the Torso Murders in 1947 but later discounted any relationship between the two cases. In 1980, new evidence implicating a former Torso Murder suspect, Jack Anderson Wilson (a.k.a. Arnold Smith), was investigated by Detective St. John in relation to Short's murder. He claimed he was close to arresting Wilson for Short's murder, but that Wilson died in a fire on February 4, 1982. The possible connection between Short's murder and the Torso Murders received renewed media attention when it was profiled on the NBC series "Unsolved Mysteries" in 1992, in which Eliot Ness biographer Oscar Fraley suggested Ness knew the identity of the killer responsible for both cases.


title: Black Dahlia

text: The February 10, 1947, murder of Jeanne French in Los Angeles was also considered by the media and detectives as possibly being connected to Short's killing. French's body was discovered in west Los Angeles on Grand View Boulevard, nude and badly beaten. Written on her stomach in lipstick was what appeared to say "Fuck You B.D.", and the letters "TEX" below. The "Herald-Express" covered the story heavily, and drew comparisons to the Short murder less than a month prior, surmising the initials "B.D." to stand for "Black Dahlia". According to historian Jon Lewis, however, the scrawling actually read "P.D.", ostensibly standing for "police department".


title: Black Dahlia

text: Crime authors such as Steve Hodel (son of George Hill Hodel) and William Rasmussen have suggested a link between the Short murder and the 1946 murder and dismemberment of six-year-old Suzanne Degnan in Chicago, Illinois. Captain Donahoe of the LAPD stated publicly that he believed the Black Dahlia and the Chicago Lipstick Murders were "likely connected". Among the evidence cited is the fact that Short's body was found on Norton Avenue, three blocks west of Degnan Boulevard, Degnan being the last name of the girl from Chicago. There were also striking similarities between the handwriting on the Degnan ransom note and that of the "Black Dahlia Avenger". Both texts used a combination of capitals and small letters (the Degnan note read in part "BuRN This FoR heR SAfTY" ), and both notes contain a similar misshapen letter P and have one word that matches exactly. Convicted serial killer William Heirens served life in prison for Degnan's murder. Initially arrested at 17 for breaking into a residence close to that of Degnan, Heirens claimed he was tortured by police, forced to confess, and made a scapegoat for the murder. After being taken from the medical infirmary at the Dixon Correctional Center on February 26, 2012, for health problems, Heirens died at the University of Illinois Medical Center on March 5, 2012, at 83.


title: Black Dahlia

text: Additionally, Steve Hodel has implicated his father, George Hodel, as Short's killer, citing his father's training as a surgeon as circumstantial evidence. In 2003, it was revealed in notes from the 1949 grand jury report that investigators had wiretapped Hodel's home, and obtained recorded conversation of him with an unidentified visitor, saying: "Supposin' I did kill the Black Dahlia. They couldn't prove it now. They can't talk to my secretary because she's dead. They thought there was something fishy. Anyway, now they may have figured it out. Killed her. Maybe I did kill my secretary."


title: Black Dahlia

text: In 1991, Janice Knowlton, a woman who was ten years old at the time of Short's murder, claimed that she witnessed her father, George Knowlton, beat Short to death with a clawhammer in the detached garage of her family's home in Westminster. She also published a book titled "Daddy was the Black Dahlia Killer" in 1995, in which she made additional claims that her father sexually molested her. The book was condemned as "trash" by Knowlton's stepsister Jolane Emerson in 2004, who stated: "She believed it, but it wasn't reality. I know, because I lived with her father for sixteen years." Additionally, Detective St. John told the "Times" that Knowlton's claims were "not consistent with the facts of the case".


title: Black Dahlia

text: John Gilmore's 1994 book "", suggests a possible connection between Short's murder and that of Georgette Bauerdorf, a socialite who was strangled to death in her West Hollywood home in 1944. Gilmore suggests that Short's employment at the Hollywood Canteen, where Bauerdorf also worked as a hostess, could be a potential connection between the two women. However, the claim that Short ever worked at the Hollywood Canteen has been disputed by others, such as the retired "Times" copyeditor Larry Harnisch (see "Rumors and factual disputes").


title: Black Dahlia

text: The 2017 book "Black Dahlia, Red Rose" by Piu Eatwell focuses on Leslie Dillon, a bellhop who was a former mortician's assistant; his associates Mark Hansen and Jeff Connors; and Sergeant Finis Brown, a lead detective who had links to Hansen and was allegedly corrupt. Eatwell posits that Short was murdered because she knew too much about the men's involvement in a scheme for robbing hotels. She further suggests that Short was killed at the Aster Motel in Los Angeles, where the owners reported finding one of their rooms "covered in blood and fecal matter" on the morning Short's body was found. The "Examiner" stated in 1949 that LA Police Chief William A. Worton denied that the Flower Street [Aster] Motel had anything to do with the case, although its rival newspaper, the "Los Angeles Herald", claimed that the murder took place there. Eatwell is working on a television documentary, and a revised edition of her book is due to be released in the autumn of 2018.


title: Black Dahlia

text: In 2000, Buz Williams, a retired detective with the Long Beach Police Department, wrote an article for the LBPD newsletter "The Rap Sheet" on Short's murder. Williams' father, Richard F. Williams, and his friend, Con Keller, were both members of LA's Gangster Squad investigating the case. Williams Sr believed that Dillon was the killer, and that when Dillon returned to his home state of Oklahoma, he was able to avoid extradition to California because his ex-wife Georgia Stevenson was second cousins with Governor Adlai Stevenson II of Illinois, who contacted the governor of Oklahoma on Dillon's behalf. Keller believed Hansen was the killer, as he had studied at a surgical school in Sweden and had thrown elaborate parties attended by prominent LAPD officials. Williams' article says that Dillon sued the LAPD for $3 million, but that the suit was dropped. Harnisch disputes this, claiming that Dillon was cleared by police after an exhaustive investigation, and that the District Attorney's files positively placed him in San Francisco when Short was killed. Harnisch claims that there was no LAPD coverup, and that Dillon did in fact receive a financial settlement from the City of Los Angeles, but has not produced concrete evidence to prove this.


title: Black Dahlia

text: Numerous details regarding Short's personal life and death have been points of public dispute. The eager involvement of both the public and press in solving her murder have been credited as factors that complicated the investigation significantly, resulting in a complex, sometimes inconsistent narrative of events. According to Anne Marie DiStefano of the "Portland Tribune", many "unsubstantiated stories" have circulated about Short over the years: "She was a prostitute, she was frigid, she was pregnant, she was a lesbian. And somehow, instead of fading away over time, the legend of the Black Dahlia just keeps getting more convoluted." Harnisch has refuted several supposed rumors and popular conceptions about Short and her murder and also disputed the validity of Gilmore's book "Severed", claiming the book is "25% mistakes, and 50% fiction". Harnisch also had examined the district attorney's files (he claimed that Steve Hodel has examined some of them pertaining to his father, along with "Times" columnist Steve Lopez) and contrary to Eatwell's claims, the files showed that Dillon was thoroughly investigated and was determined to have been in San Francisco when Short was killed. Harnisch speculated that Eatwell either did not find these files or she chose to ignore them.


title: Black Dahlia

text: A number of people, none of whom knew Short, contacted police and the newspapers and claimed to have seen her during her so-called "missing week", between her January 9 disappearance and the discovery of her body, on January 15. Police and DA investigators ruled out each alleged sighting; in some cases, those interviewed were identifying other women whom they had mistaken for Short. Short's whereabouts in the days leading up to her murder and the discovery of her body are unknown.


title: Black Dahlia

text: After the discovery of Short's body, numerous Los Angeles newspapers printed headlines claiming she had been tortured leading up to her death. This was denied by law enforcement at the time, but they allowed the claims to circulate so as to keep Short's actual cause of death a secret from the public. Some sources, such as Oliver Cyriax's "Crime: An Encyclopedia" (1993), state that Short's body was covered in cigarette burns inflicted on her while she was still alive, though there is no indication of this in her official autopsy report.


title: Black Dahlia

text: In "Severed", Gilmore states that the coroner who performed Short's autopsy suggested in conversation that she had been forced to consume feces based on his findings when examining the contents of her stomach. This claim has been denied by Harnisch and is also not indicated in Short's official autopsy, though it has been reprinted in several print and online media.


title: Black Dahlia

text: According to newspaper reports shortly after the murder, Short received the nickname "Black Dahlia" from staff and patrons at a Long Beach drugstore in mid-1946 as wordplay on the film "The Blue Dahlia" (1946). Other popularly-circulated rumors claim that the media crafted the name because Short adorned her hair with dahlias. According to the FBI official website, she received the first part of the nickname from the press "for her rumored penchant for sheer black clothes".


title: Black Dahlia

text: However, reports by DA investigators state that the nickname was invented by newspaper reporters covering her murder; "Herald-Express" reporter Bevo Means, who interviewed Short's acquaintances at the drugstore, has been credited with first using the "Black Dahlia" name, though reporters Underwood and Jack Smith have been alternatively named as its creators. While some sources claim that Short was referred to or went by the name during her life, others dispute this. Both Gilmore and Harnisch agree that the name originated during Short's lifetime and was not a creation of the press: Harnisch states that it was in fact a nickname she earned from the staff of the Long Beach drugstore she frequented; in "Severed", Gilmore names an A.L. Landers as the proprietor of the drugstore, though he does not provide the store's name. Prior to the circulation of the "Black Dahlia" name, Short's killing had been dubbed the "Werewolf Murder" by the "Herald-Express" because of the brutal nature of the crime.


title: Black Dahlia

text: Many true crime books claim that Short lived in or visited Los Angeles at various times in the mid-1940s, including Gilmore's "Severed", which claims she worked at the Hollywood Canteen. This is disputed by Harnisch, who states that Short did not, in fact, live in Los Angeles until after the canteen's closing in 1945. Although some of her acquaintances and several authors and journalists described Short as a prostitute or call girl during her time in Los Angeles, according to Harnisch, the contemporaneous grand jury proved that there was no existing evidence that she was ever a prostitute. Harnisch claims that the rumor regarding Short's history as a prostitute originates from John Gregory Dunne's 1977 novel "True Confessions", which is based in part on the crime.


title: Black Dahlia

text: Another widely circulated rumor (sometimes used to counter claims that Short was a prostitute) holds that Short was unable to have sexual intercourse because of a congenital defect that resulted in gonadal dysgenesis, also known as "infantile genitalia". Los Angeles County district attorney's files state that the investigators had questioned three men with whom Short had engaged in sex, including a Chicago police officer who was a suspect in the case; FBI files on the case also contain a statement from one of Short's alleged lovers. Short's autopsy itself, which was reprinted in full in Michael Newton's 2009 book "The Encyclopedia of Unsolved Crimes", notes that her uterus was "small"; however, no other information in the autopsy is provided that would suggest her reproductive organs were anything other than anatomically normal. The autopsy also states that Short was not and had never been pregnant, contrary to what had been claimed prior to and following her death.


title: Black Dahlia

text: Another rumor—that Short was a lesbian—has often circulated; according to Gilmore, this rumor began after Bevo Means of the "Herald-Express" was told by the deputy coroner that Short "wasn't having sex with men" owing to her allegedly "small" genitalia. Means took this to mean that Short had sex with women, and both he and reporter Sid Hughes began fruitlessly investigating gay bars in Los Angeles for further information.


title: Black Dahlia

text: Short is interred at the Mountain View Cemetery in Oakland. After her younger sisters had grown up and married, their mother, Phoebe, moved to Oakland to be near her daughter's grave. She finally returned to the East Coast in the 1970s, where she lived into her 90s. On February 2, 1947, just two weeks after Short's murder, Republican state assemblyman C. Don Field was prompted by the case to introduce a bill calling for the formation of a sex offender registry; the state of California would become the first U.S. state to make the registration of sex offenders mandatory.


title: Black Dahlia

text: Short's murder has been described as one of the most brutal and culturally enduring crimes in American history, and "Time" magazine listed it as one of the most infamous unsolved cases in the world.


title: Black Dahlia

text: Short's life and death have been the basis of numerous books, television shows, and films, both fictionalized and non-fiction. Among the most famous fictional accounts of Short's death is James Ellroy's 1987 novel "The Black Dahlia", which, in addition to the murder, explored "the larger fields of politics, crime, corruption, and paranoia in post-war Los Angeles", according to cultural critic David M. Fine. Ellroy's novel was adapted into a 2006 film of the same name by director Brian De Palma: Short was played by actress Mia Kirshner. Both Ellroy's novel and its film adaptation bear little relation to the facts of the case.


title: Black Dahlia

text: Short was also portrayed in heavily fictionalized accounts by Lucie Arnaz in the 1975 television film "Who Is the Black Dahlia?", by Jessica Nelson in Season Four, Episode 13 of "Hunter", and by Mena Suvari in the series "American Horror Story" in 2011, featuring Short in the plot line of the episode "Spooky Little Girl", and again in 2018 with "Return to Murder House".


title: Arthropod

text: Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, often mineralised with calcium carbonate. The arthropod body plan consists of segments, each with a pair of appendages. Arthropods are bilaterally symmetrical and their body possesses an external skeleton. In order to keep growing, they must go through stages of moulting, a process by which they shed their exoskeleton to reveal a new one. Some species have wings. They are an extremely diverse group, with up to 10 million species.


title: Arthropod

text: The haemocoel, an arthropod's internal cavity, through which its haemolymph – analogue of blood – circulates, accommodates its interior organs; it has an open circulatory system. Like their exteriors, the internal organs of arthropods are generally built of repeated segments. Their nervous system is "ladder-like", with paired ventral nerve cords running through all segments and forming paired ganglia in each segment. Their heads are formed by fusion of varying numbers of segments, and their brains are formed by fusion of the ganglia of these segments and encircle the esophagus. The respiratory and excretory systems of arthropods vary, depending as much on their environment as on the subphylum to which they belong.


title: Arthropod

text: Arthropods use combinations of compound eyes and pigment-pit ocelli for vision. In most species, the ocelli can only detect the direction from which light is coming, and the compound eyes are the main source of information, but the main eyes of spiders are ocelli that can form images and, in a few cases, can swivel to track prey. Arthropods also have a wide range of chemical and mechanical sensors, mostly based on modifications of the many bristles known as setae that project through their cuticles. Similarly, their reproduction and development are varied; all terrestrial species use internal fertilization, but this is sometimes by indirect transfer of the sperm via an appendage or the ground, rather than by direct injection. Aquatic species use either internal or external fertilization. Almost all arthropods lay eggs, but many species give birth to live young after the eggs have hatched inside the mother, and a few are genuinely viviparous, such as aphids. Arthropod hatchlings vary from miniature adults to grubs and caterpillars that lack jointed limbs and eventually undergo a total metamorphosis to produce the adult form. The level of maternal care for hatchlings varies from nonexistent to the prolonged care provided by social insects.


title: Arthropod

text: The evolutionary ancestry of arthropods dates back to the Cambrian period. The group is generally regarded as monophyletic, and many analyses support the placement of arthropods with cycloneuralians (or their constituent clades) in a superphylum Ecdysozoa. Overall, however, the basal relationships of animals are not yet well resolved. Likewise, the relationships between various arthropod groups are still actively debated. Today, Arthropods contribute to the human food supply both directly as food, and more importantly, indirectly as pollinators of crops. Some species are known to spread severe disease to humans, livestock, and crops.


title: Arthropod

text: The word "arthropod" comes from the Greek "árthron", "joint", and "pous" (gen. "podos (ποδός)"), i.e. "foot" or "leg", which together mean "jointed leg". The designation "Arthropoda" was coined in 1848 by the German physiologist and zoologist Karl Theodor Ernst von Siebold (1804–1885).


title: Arthropod

text: In common parlance, terrestrial arthropods are often called bugs. The term is also occasionally extended to colloquial names for freshwater or marine crustaceans (e.g. Balmain bug, Moreton Bay bug, mudbug) and used by physicians and bacteriologists for disease-causing germs (e.g. superbugs), but entomologists reserve this term for a narrow category of "true bugs", insects of the order Hemiptera (which does not include ants, bees, beetles, butterflies or moths).


title: Arthropod

text: Arthropods are invertebrates with segmented bodies and jointed limbs. The exoskeleton or cuticles consists of chitin, a polymer of glucosamine. The cuticle of many crustaceans, beetle mites, and millipedes (except for bristly millipedes) is also biomineralized with calcium carbonate. Calcification of the endosternite, an internal structure used for muscle attachments, also occur in some opiliones.


title: Arthropod

text: Estimates of the number of arthropod species vary between 1,170,000 and 5 to 10 million and account for over 80 percent of all known living animal species. The number of species remains difficult to determine. This is due to the census modeling assumptions projected onto other regions in order to scale up from counts at specific locations applied to the whole world. A study in 1992 estimated that there were 500,000 species of animals and plants in Costa Rica alone, of which 365,000 were arthropods.


title: Arthropod

text: They are important members of marine, freshwater, land and air ecosystems, and are one of only two major animal groups that have adapted to life in dry environments; the other is amniotes, whose living members are reptiles, birds and mammals. One arthropod sub-group, insects, is the most species-rich member of all ecological guilds in land and freshwater environments. The lightest insects weigh less than 25 micrograms (millionths of a gram), while the heaviest weigh over . Some living malacostracans are much larger; for example, the legs of the Japanese spider crab may span up to , with the heaviest of all living arthropods being the American lobster, topping out at over 20 kg (44 lbs).


title: Arthropod

text: The embryos of all arthropods are segmented, built from a series of repeated modules. The last common ancestor of living arthropods probably consisted of a series of undifferentiated segments, each with a pair of appendages that functioned as limbs. However, all known living and fossil arthropods have grouped segments into tagmata in which segments and their limbs are specialized in various ways.


title: Arthropod

text: The three-part appearance of many insect bodies and the two-part appearance of spiders is a result of this grouping; in fact there are no external signs of segmentation in mites. Arthropods also have two body elements that are not part of this serially repeated pattern of segments, an ocular somite at the front, where the mouth and eyes originated, and a telson at the rear, behind the anus.


title: Arthropod

text: Originally it seems that each appendage-bearing segment had two separate pairs of appendages: an upper, unsegmented exite and a lower, segmented endopod. These would later fuse into a single pair of biramous appendages united by a basal segment (protopod or basipod), with the upper branch acting as a gill while the lower branch was used for locomotion. The appendages of most crustaceans and some extinct taxa such as trilobites have another segmented branch known as exopods, but whether these structures have a single origin remain controversial. In some segments of all known arthropods the appendages have been modified, for example to form gills, mouth-parts, antennae for collecting information, or claws for grasping; arthropods are "like Swiss Army knives, each equipped with a unique set of specialized tools." In many arthropods, appendages have vanished from some regions of the body; it is particularly common for abdominal appendages to have disappeared or be highly modified.


title: Arthropod

text: The most conspicuous specialization of segments is in the head. The four major groups of arthropods – Chelicerata (sea spiders, horseshoe crabs and arachnids), Myriapoda (symphylan, pauropods, millipedes and centipedes), Crustacea (oligostracans, copepods, malacostracans, branchiopods, hexapods, etc.), and the extinct Trilobita – have heads formed of various combinations of segments, with appendages that are missing or specialized in different ways. Despite myriapods and hexapods both having similar head combinations, hexapods are deeply nested within crustacea while myriapods are not, so these traits are believed to have evolved separately. In addition, some extinct arthropods, such as "Marrella", belong to none of these groups, as their heads are formed by their own particular combinations of segments and specialized appendages.


title: Arthropod

text: Working out the evolutionary stages by which all these different combinations could have appeared is so difficult that it has long been known as "the arthropod head problem". In 1960, R. E. Snodgrass even hoped it would not be solved, as he found trying to work out solutions to be fun.


title: Arthropod

text: Arthropod exoskeletons are made of cuticle, a non-cellular material secreted by the epidermis. Their cuticles vary in the details of their structure, but generally consist of three main layers: the epicuticle, a thin outer waxy coat that moisture-proofs the other layers and gives them some protection; the exocuticle, which consists of chitin and chemically hardened proteins; and the endocuticle, which consists of chitin and unhardened proteins. The exocuticle and endocuticle together are known as the procuticle. Each body segment and limb section is encased in hardened cuticle. The joints between body segments and between limb sections are covered by flexible cuticle.


title: Arthropod

text: The exoskeletons of most aquatic crustaceans are biomineralized with calcium carbonate extracted from the water. Some terrestrial crustaceans have developed means of storing the mineral, since on land they cannot rely on a steady supply of dissolved calcium carbonate. Biomineralization generally affects the exocuticle and the outer part of the endocuticle. Two recent hypotheses about the evolution of biomineralization in arthropods and other groups of animals propose that it provides tougher defensive armor, and that it allows animals to grow larger and stronger by providing more rigid skeletons; and in either case a mineral-organic composite exoskeleton is cheaper to build than an all-organic one of comparable strength.


title: Arthropod

text: The cuticle may have setae (bristles) growing from special cells in the epidermis. Setae are as varied in form and function as appendages. For example, they are often used as sensors to detect air or water currents, or contact with objects; aquatic arthropods use feather-like setae to increase the surface area of swimming appendages and to filter food particles out of water; aquatic insects, which are air-breathers, use thick felt-like coats of setae to trap air, extending the time they can spend under water; heavy, rigid setae serve as defensive spines.


title: Arthropod

text: Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, some still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors; for example, all spiders extend their legs hydraulically and can generate pressures up to eight times their resting level.


title: Arthropod

text: The exoskeleton cannot stretch and thus restricts growth. Arthropods, therefore, replace their exoskeletons by undergoing ecdysis (moulting), or shedding the old exoskeleton after growing a new one that is not yet hardened. Moulting cycles run nearly continuously until an arthropod reaches full size.


title: Arthropod

text: The developmental stages between each moult (ecdysis) until sexual maturity is reached is called an instar. Differences between instars can often be seen in altered body proportions, colors, patterns, changes in the number of body segments or head width. After moulting, i.e. shedding their exoskeleton, the juvenile arthropods continue in their life cycle until they either pupate or moult again.


title: Arthropod

text: In the initial phase of moulting, the animal stops feeding and its epidermis releases moulting fluid, a mixture of enzymes that digests the endocuticle and thus detaches the old cuticle. This phase begins when the epidermis has secreted a new epicuticle to protect it from the enzymes, and the epidermis secretes the new exocuticle while the old cuticle is detaching. When this stage is complete, the animal makes its body swell by taking in a large quantity of water or air, and this makes the old cuticle split along predefined weaknesses where the old exocuticle was thinnest. It commonly takes several minutes for the animal to struggle out of the old cuticle. At this point, the new one is wrinkled and so soft that the animal cannot support itself and finds it very difficult to move, and the new endocuticle has not yet formed. The animal continues to pump itself up to stretch the new cuticle as much as possible, then hardens the new exocuticle and eliminates the excess air or water. By the end of this phase, the new endocuticle has formed. Many arthropods then eat the discarded cuticle to reclaim its materials.


title: Arthropod

text: Because arthropods are unprotected and nearly immobilized until the new cuticle has hardened, they are in danger both of being trapped in the old cuticle and of being attacked by predators. Moulting may be responsible for 80 to 90% of all arthropod deaths.


title: Arthropod

text: Arthropod bodies are also segmented internally, and the nervous, muscular, circulatory, and excretory systems have repeated components. Arthropods come from a lineage of animals that have a coelom, a membrane-lined cavity between the gut and the body wall that accommodates the internal organs. The strong, segmented limbs of arthropods eliminate the need for one of the coelom's main ancestral functions, as a hydrostatic skeleton, which muscles compress in order to change the animal's shape and thus enable it to move. Hence the coelom of the arthropod is reduced to small areas around the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows.


title: Arthropod

text: Arthropods have open circulatory systems, although most have a few short, open-ended arteries. In chelicerates and crustaceans, the blood carries oxygen to the tissues, while hexapods use a separate system of tracheae. Many crustaceans, but few chelicerates and tracheates, use respiratory pigments to assist oxygen transport. The most common respiratory pigment in arthropods is copper-based hemocyanin; this is used by many crustaceans and a few centipedes. A few crustaceans and insects use iron-based hemoglobin, the respiratory pigment used by vertebrates. As with other invertebrates, the respiratory pigments of those arthropods that have them are generally dissolved in the blood and rarely enclosed in corpuscles as they are in vertebrates.


title: Arthropod

text: The heart is typically a muscular tube that runs just under the back and for most of the length of the hemocoel. It contracts in ripples that run from rear to front, pushing blood forwards. Sections not being squeezed by the heart muscle are expanded either by elastic ligaments or by small muscles, in either case connecting the heart to the body wall. Along the heart run a series of paired ostia, non-return valves that allow blood to enter the heart but prevent it from leaving before it reaches the front.


title: Arthropod

text: Arthropods have a wide variety of respiratory systems. Small species often do not have any, since their high ratio of surface area to volume enables simple diffusion through the body surface to supply enough oxygen. Crustacea usually have gills that are modified appendages. Many arachnids have book lungs. Tracheae, systems of branching tunnels that run from the openings in the body walls, deliver oxygen directly to individual cells in many insects, myriapods and arachnids.


title: Arthropod

text: Living arthropods have paired main nerve cords running along their bodies below the gut, and in each segment the cords form a pair of ganglia from which sensory and motor nerves run to other parts of the segment. Although the pairs of ganglia in each segment often appear physically fused, they are connected by commissures (relatively large bundles of nerves), which give arthropod nervous systems a characteristic "ladder-like" appearance. The brain is in the head, encircling and mainly "above" the esophagus. It consists of the fused ganglia of the acron and one or two of the foremost segments that form the head – a total of three pairs of ganglia in most arthropods, but only two in chelicerates, which do not have antennae or the ganglion connected to them. The ganglia of other head segments are often close to the brain and function as part of it. In insects these other head ganglia combine into a pair of subesophageal ganglia, under and behind the esophagus. Spiders take this process a step further, as "all" the segmental ganglia are incorporated into the subesophageal ganglia, which occupy most of the space in the cephalothorax (front "super-segment").


title: Arthropod

text: There are two different types of arthropod excretory systems. In aquatic arthropods, the end-product of biochemical reactions that metabolise nitrogen is ammonia, which is so toxic that it needs to be diluted as much as possible with water. The ammonia is then eliminated via any permeable membrane, mainly through the gills. All crustaceans use this system, and its high consumption of water may be responsible for the relative lack of success of crustaceans as land animals. Various groups of terrestrial arthropods have independently developed a different system: the end-product of nitrogen metabolism is uric acid, which can be excreted as dry material; the Malpighian tubule system filters the uric acid and other nitrogenous waste out of the blood in the hemocoel, and dumps these materials into the hindgut, from which they are expelled as feces. Most aquatic arthropods and some terrestrial ones also have organs called nephridia ("little kidneys"), which extract other wastes for excretion as urine.


title: Arthropod

text: The stiff cuticles of arthropods would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae. Pressure sensors often take the form of membranes that function as eardrums, but are connected directly to nerves rather than to auditory ossicles. The antennae of most hexapods include sensor packages that monitor humidity, moisture and temperature.


title: Arthropod

text: Most arthropods lack balance and acceleration sensors, and rely on their eyes to tell them which way is up. The self-righting behavior of cockroaches is triggered when pressure sensors on the underside of the feet report no pressure. However, many malacostracan crustaceans have statocysts, which provide the same sort of information as the balance and motion sensors of the vertebrate inner ear.


title: Arthropod

text: The proprioceptors of arthropods, sensors that report the force exerted by muscles and the degree of bending in the body and joints, are well understood. However, little is known about what other internal sensors arthropods may have.


title: Arthropod

text: Most arthropods have sophisticated visual systems that include one or more usually both of compound eyes and pigment-cup ocelli ("little eyes"). In most cases ocelli are only capable of detecting the direction from which light is coming, using the shadow cast by the walls of the cup. However, the main eyes of spiders are pigment-cup ocelli that are capable of forming images, and those of jumping spiders can rotate to track prey.


title: Arthropod

text: Compound eyes consist of fifteen to several thousand independent ommatidia, columns that are usually hexagonal in cross section. Each ommatidium is an independent sensor, with its own light-sensitive cells and often with its own lens and cornea. Compound eyes have a wide field of view, and can detect fast movement and, in some cases, the polarization of light. On the other hand, the relatively large size of ommatidia makes the images rather coarse, and compound eyes are shorter-sighted than those of birds and mammals – although this is not a severe disadvantage, as objects and events within are most important to most arthropods. Several arthropods have color vision, and that of some insects has been studied in detail; for example, the ommatidia of bees contain receptors for both green and ultra-violet.


title: Arthropod

text: A few arthropods, such as barnacles, are hermaphroditic, that is, each can have the organs of both sexes. However, individuals of most species remain of one sex their entire lives. A few species of insects and crustaceans can reproduce by parthenogenesis, especially if conditions favor a "population explosion". However, most arthropods rely on sexual reproduction, and parthenogenetic species often revert to sexual reproduction when conditions become less favorable. The ability to undergo meiosis is widespread among arthropods including both those that reproduce sexually and those that reproduce parthenogenetically. Although meiosis is a major characteristic of arthropods, understanding of its fundamental adaptive benefit has long been regarded as an unresolved problem, that appears to have remained unsettled.


title: Arthropod

text: Aquatic arthropods may breed by external fertilization, as for example horseshoe crabs do, or by internal fertilization, where the ova remain in the female's body and the sperm must somehow be inserted. All known terrestrial arthropods use internal fertilization. Opiliones (harvestmen), millipedes, and some crustaceans use modified appendages such as gonopods or penises to transfer the sperm directly to the female. However, most male terrestrial arthropods produce spermatophores, waterproof packets of sperm, which the females take into their bodies. A few such species rely on females to find spermatophores that have already been deposited on the ground, but in most cases males only deposit spermatophores when complex courtship rituals look likely to be successful.


title: Arthropod

text: Most arthropods lay eggs, but scorpions are ovoviparous: they produce live young after the eggs have hatched inside the mother, and are noted for prolonged maternal care. Newly born arthropods have diverse forms, and insects alone cover the range of extremes. Some hatch as apparently miniature adults (direct development), and in some cases, such as silverfish, the hatchlings do not feed and may be helpless until after their first moult. Many insects hatch as grubs or caterpillars, which do not have segmented limbs or hardened cuticles, and metamorphose into adult forms by entering an inactive phase in which the larval tissues are broken down and re-used to build the adult body. Dragonfly larvae have the typical cuticles and jointed limbs of arthropods but are flightless water-breathers with extendable jaws. Crustaceans commonly hatch as tiny nauplius larvae that have only three segments and pairs of appendages.


title: Arthropod

text: Based on the distribution of shared plesiomorphic features in extant and fossil taxa, the last common ancestor of all arthropods is inferred to have been as a modular organism with each module covered by its own sclerite (armor plate) and bearing a pair of biramous limbs. However, whether the ancestral limb was uniramous or biramous is far from a settled debate.


title: Arthropod

text: This Ur-arthropod had a ventral mouth, pre-oral antennae and dorsal eyes at the front of the body. It was assumed to have been a non-discriminatory sediment feeder, processing whatever sediment came its way for food, but fossil findings hint that the last common ancestor of both arthropods and priapulida shared the same specialized mouth apparatus; a circular mouth with rings of teeth used for capturing animal prey.


title: Arthropod

text: It has been proposed that the Ediacaran animals "Parvancorina" and "Spriggina", from around , were arthropods, but later study shows that their affinities of being origin of arthropods are not reliable. Small arthropods with bivalve-like shells have been found in Early Cambrian fossil beds dating in China and Australia. The earliest Cambrian trilobite fossils are about 530 million years old, but the class was already quite diverse and worldwide, suggesting that they had been around for quite some time. In the Maotianshan shales, which date to between 530 and 520 million years ago, fossils of arthropods such as "Kylinxia" and "Erratus" have been found that seem to show a transitional split between lobopodia and other more primitive stem arthropods. Re-examination in the 1970s of the Burgess Shale fossils from about identified many arthropods, some of which could not be assigned to any of the well-known groups, and thus intensified the debate about the Cambrian explosion. A fossil of "Marrella" from the Burgess Shale has provided the earliest clear evidence of moulting.


title: Arthropod

text: The earliest fossil crustaceans date from about in the Cambrian, and fossil shrimp from about apparently formed a tight-knit procession across the seabed. Crustacean fossils are common from the Ordovician period onwards. They have remained almost entirely aquatic, possibly because they never developed excretory systems that conserve water. In 2020 scientists announced the discovery of "Kylinxia", a five-eyed ~5 cm long shrimp-like animal living 518 Mya that – with multiple distinctive features – appears to be a key ‘missing link’ of the evolution from "Anomalocaris" to true arthropods and could be at the evolutionary root of true arthropods.


title: Arthropod

text: Arthropods provide the earliest identifiable fossils of land animals, from about in the Late Silurian, and terrestrial tracks from about appear to have been made by arthropods. Arthropods possessed attributes that were easy coopted for life on land; their existing jointed exoskeletons provided protection against desiccation, support against gravity and a means of locomotion that was not dependent on water. Around the same time the aquatic, scorpion-like eurypterids became the largest ever arthropods, some as long as .


title: Arthropod

text: The oldest known arachnid is the trigonotarbid "Palaeotarbus jerami", from about in the Silurian period. "Attercopus fimbriunguis", from in the Devonian period, bears the earliest known silk-producing spigots, but its lack of spinnerets means it was not one of the true spiders, which first appear in the Late Carboniferous over . The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families. Fossils of aquatic scorpions with gills appear in the Silurian and Devonian periods, and the earliest fossil of an air-breathing scorpion with book lungs dates from the Early Carboniferous period.


title: Arthropod

text: The oldest possible insect fossil is the Devonian "Rhyniognatha hirsti", dated at , but its mandibles are of a type found only in winged insects, which suggests that the earliest insects appeared in the Silurian period, although later study shows possibility that "Rhyniognatha" can be myriapod, not an insect. The Mazon Creek lagerstätten from the Late Carboniferous, about , include about 200 species, some gigantic by modern standards, and indicate that insects had occupied their main modern ecological niches as herbivores, detritivores and insectivores. Social termites and ants first appear in the Early Cretaceous, and advanced social bees have been found in Late Cretaceous rocks but did not become abundant until the Middle Cenozoic.


title: Arthropod

text: From 1952 to 1977, zoologist Sidnie Manton and others argued that arthropods are polyphyletic, in other words, that they do not share a common ancestor that was itself an arthropod. Instead, they proposed that three separate groups of "arthropods" evolved separately from common worm-like ancestors: the chelicerates, including spiders and scorpions; the crustaceans; and the uniramia, consisting of onychophorans, myriapods and hexapods. These arguments usually bypassed trilobites, as the evolutionary relationships of this class were unclear. Proponents of polyphyly argued the following: that the similarities between these groups are the results of convergent evolution, as natural consequences of having rigid, segmented exoskeletons; that the three groups use different chemical means of hardening the cuticle; that there were significant differences in the construction of their compound eyes; that it is hard to see how such different configurations of segments and appendages in the head could have evolved from the same ancestor; and that crustaceans have biramous limbs with separate gill and leg branches, while the other two groups have uniramous limbs in which the single branch serves as a leg.


title: Arthropod

text: Further analysis and discoveries in the 1990s reversed this view, and led to acceptance that arthropods are monophyletic, in other words they are inferred to share a common ancestor that was itself an arthropod. For example, Graham Budd's analyses of "Kerygmachela" in 1993 and of "Opabinia" in 1996 convinced him that these animals were similar to onychophorans and to various Early Cambrian "lobopods", and he presented an "evolutionary family tree" that showed these as "aunts" and "cousins" of all arthropods. These changes made the scope of the term "arthropod" unclear, and Claus Nielsen proposed that the wider group should be labelled "Panarthropoda" ("all the arthropods") while the animals with jointed limbs and hardened cuticles should be called "Euarthropoda" ("true arthropods").


title: Arthropod

text: A contrary view was presented in 2003, when Jan Bergström and Xian-Guang Hou argued that, if arthropods were a "sister-group" to any of the anomalocarids, they must have lost and then re-evolved features that were well-developed in the anomalocarids. The earliest known arthropods ate mud in order to extract food particles from it, and possessed variable numbers of segments with unspecialized appendages that functioned as both gills and legs. Anomalocarids were, by the standards of the time, huge and sophisticated predators with specialized mouths and grasping appendages, fixed numbers of segments some of which were specialized, tail fins, and gills that were very different from those of arthropods. In 2006, they suggested that arthropods were more closely related to lobopods and tardigrades than to anomalocarids. In 2014, research indicated that tardigrades were more closely related to arthropods than velvet worms.


title: Arthropod

text: Higher up the "family tree", the Annelida have traditionally been considered the closest relatives of the Panarthropoda, since both groups have segmented bodies, and the combination of these groups was labelled Articulata. There had been competing proposals that arthropods were closely related to other groups such as nematodes, priapulids and tardigrades, but these remained minority views because it was difficult to specify in detail the relationships between these groups.


title: Arthropod

text: In the 1990s, molecular phylogenetic analyses of DNA sequences produced a coherent scheme showing arthropods as members of a superphylum labelled Ecdysozoa ("animals that moult"), which contained nematodes, priapulids and tardigrades but excluded annelids. This was backed up by studies of the anatomy and development of these animals, which showed that many of the features that supported the Articulata hypothesis showed significant differences between annelids and the earliest Panarthropods in their details, and some were hardly present at all in arthropods. This hypothesis groups annelids with molluscs and brachiopods in another superphylum, Lophotrochozoa.


title: Arthropod

text: If the Ecdysozoa hypothesis is correct, then segmentation of arthropods and annelids either has evolved convergently or has been inherited from a much older ancestor and subsequently lost in several other lineages, such as the non-arthropod members of the Ecdysozoa.


title: Arthropod

text: Modern interpretations of the basal, extinct stem-group of Arthropoda recognised the following groups, from most basal to most crownward:


title: Arthropod

text: The Deuteropoda is a recently established clade uniting the crown-group (living) arthropods with these possible "upper stem-group" fossils taxa. The clade is defined by important changes to the structure of the head region such as the appearance of a differentiated deutocerebral appendage pair.


title: Arthropod

text: However, recent analyses since late 2010s also show that these "upper stem-groups" might be inside the crown-group: isoxyids might nested with the crown-group itself, Megacheira have been recovered as more closely related to Chelicerates, some bivalved forms such as Hymenocarina are consistently shown to be mandibulates, and similarly Fuxianhuiida might also be mandibulates as well.


title: Arthropod

text: The following cladogram shows the probable relationships between crown-group Arthropoda and stem-group Arthropoda according to O’Flynn et al. 2022, including two new fossils found to be the most early branches of Deuteropoda (the "upper stem-groups" in previous studies are marked in asterisk, living groups are marked in bold):


title: Arthropod

text: Note that the subphylum Artiopoda, containing the trilobites, is closer to mandibulates than to chelicerates in the cladogram above, but older analyses place them as the sister group of chelicerates united under the clade Arachnomorpha.


title: Arthropod

text: The following cladogram shows the internal relationships between all the living classes of arthropods as of late 2010s, as well as the estimated timing for some of the clades:


title: Arthropod

text: Aside from these major groups, a number of fossil forms, mostly from the early Cambrian period, are difficult to place taxonomically, either from lack of obvious affinity to any of the main groups or from clear affinity to several of them. "Marrella" was the first one to be recognized as significantly different from the well-known groups.


title: Arthropod

text: The phylogeny of the major extant arthropod groups has been an area of considerable interest and dispute. Recent studies strongly suggest that Crustacea, as traditionally defined, is paraphyletic, with Hexapoda having evolved from within it, so that Crustacea and Hexapoda form a clade, Pancrustacea. The position of Myriapoda, Chelicerata and Pancrustacea remains unclear . In some studies, Myriapoda is grouped with Chelicerata (forming Myriochelata); in other studies, Myriapoda is grouped with Pancrustacea (forming Mandibulata), or Myriapoda may be sister to Chelicerata plus Pancrustacea.


title: Arthropod

text: The placement of the extinct trilobites is also a frequent subject of dispute. One of the newer hypotheses is that the chelicerae have originated from the same pair of appendages that evolved into antennae in the ancestors of Mandibulata, which would place trilobites, which had antennae, closer to Mandibulata than Chelicerata.


title: Arthropod

text: Since the International Code of Zoological Nomenclature recognises no priority above the rank of family, many of the higher-level groups can be referred to by a variety of different names.


title: Arthropod

text: Crustaceans such as crabs, lobsters, crayfish, shrimp, and prawns have long been part of human cuisine, and are now raised commercially. Insects and their grubs are at least as nutritious as meat, and are eaten both raw and cooked in many cultures, though not most European, Hindu, and Islamic cultures. Cooked tarantulas are considered a delicacy in Cambodia, and by the Piaroa Indians of southern Venezuela, after the highly irritant hairs – the spider's main defense system – are removed. Humans also unintentionally eat arthropods in other foods, and food safety regulations lay down acceptable contamination levels for different kinds of food material. The intentional cultivation of arthropods and other small animals for human food, referred to as minilivestock, is now emerging in animal husbandry as an ecologically sound concept. Commercial butterfly breeding provides Lepidoptera stock to butterfly conservatories, educational exhibits, schools, research facilities, and cultural events.


title: Arthropod

text: However, the greatest contribution of arthropods to human food supply is by pollination: a 2008 study examined the 100 crops that FAO lists as grown for food, and estimated pollination's economic value as €153 billion, or 9.5 per cent of the value of world agricultural production used for human food in 2005. Besides pollinating, bees produce honey, which is the basis of a rapidly growing industry and international trade.


title: Arthropod

text: The red dye cochineal, produced from a Central American species of insect, was economically important to the Aztecs and Mayans. While the region was under Spanish control, it became Mexico's second most-lucrative export, and is now regaining some of the ground it lost to synthetic competitors. Shellac, a resin secreted by a species of insect native to southern Asia, was historically used in great quantities for many applications in which it has mostly been replaced by synthetic resins, but it is still used in woodworking and as a food additive. The blood of horseshoe crabs contains a clotting agent, Limulus Amebocyte Lysate, which is now used to test that antibiotics and kidney machines are free of dangerous bacteria, and to detect spinal meningitis and some cancers. Forensic entomology uses evidence provided by arthropods to establish the time and sometimes the place of death of a human, and in some cases the cause. Recently insects have also gained attention as potential sources of drugs and other medicinal substances.


title: Arthropod

text: The relative simplicity of the arthropods' body plan, allowing them to move on a variety of surfaces both on land and in water, have made them useful as models for robotics. The redundancy provided by segments allows arthropods and biomimetic robots to move normally even with damaged or lost appendages.


title: Arthropod

text: Although arthropods are the most numerous phylum on Earth, and thousands of arthropod species are venomous, they inflict relatively few serious bites and stings on humans. Far more serious are the effects on humans of diseases like malaria carried by blood-sucking insects. Other blood-sucking insects infect livestock with diseases that kill many animals and greatly reduce the usefulness of others. Ticks can cause tick paralysis and several parasite-borne diseases in humans. A few of the closely related mites also infest humans, causing intense itching, and others cause allergic diseases, including hay fever, asthma, and eczema.


title: Arthropod

text: Many species of arthropods, principally insects but also mites, are agricultural and forest pests. The mite "Varroa destructor" has become the largest single problem faced by beekeepers worldwide. Efforts to control arthropod pests by large-scale use of pesticides have caused long-term effects on human health and on biodiversity. Increasing arthropod resistance to pesticides has led to the development of integrated pest management using a wide range of measures including biological control. Predatory mites may be useful in controlling some mite pests.


title: Arthropod

text: Even amongst arthropods usually thought of as obligate predators, floral food sources (nectar and to a lesser degree pollen) are often useful adjunct sources. It was noticed in one study that adult "Adalia bipunctata" (predator and common biocontrol of "Ephestia kuehniella") could survive on flowers but never completed the life cycle, so a meta-analysis was done to find such an overall trend in previously published data, if it existed. In some cases floral resources are outright necessary. Overall, floral resources (and an imitation, i.e. sugar water) increase longevity and fecundity, meaning even predatory population numbers can depend on non-prey food abundance. Thus biocontrol success may surprisingly depend on nearby flowers.


title: Boeing AH-64 Apache

text: The Boeing AH-64 Apache () is an American twin-turboshaft attack helicopter with a tailwheel-type landing gear arrangement and a tandem cockpit for a crew of two. It features a nose-mounted sensor suite for target acquisition and night vision systems. It is armed with a M230 chain gun carried between the main landing gear, under the aircraft's forward fuselage, and four hardpoints mounted on stub-wing pylons for carrying armament and stores, typically a mixture of AGM-114 Hellfire missiles and Hydra 70 rocket pods. The AH-64 has significant systems redundancy to improve combat survivability.


title: Boeing AH-64 Apache

text: The Apache began as the "Model 77" developed by Hughes Helicopters for the United States Army's Advanced Attack Helicopter program to replace the AH-1 Cobra. The prototype YAH-64 was first flown on 30 September 1975. The U.S. Army selected the YAH-64 over the Bell YAH-63 in 1976, and later approved full production in 1982. After purchasing Hughes Helicopters in 1984, McDonnell Douglas continued AH-64 production and development. The helicopter was introduced to U.S. Army service in April 1986. The advanced AH-64D Apache Longbow was delivered to the Army in March 1997. Production has been continued by Boeing Defense, Space & Security, with over 2,400 AH-64s being produced by 2020.


title: Boeing AH-64 Apache

text: The U.S. Army is the primary operator of the AH-64. It has also become the primary attack helicopter of multiple nations, including Greece, Japan, Israel, the Netherlands, Singapore, and the United Arab Emirates. It has been built under license in the United Kingdom as the AgustaWestland Apache. American AH-64s have served in conflicts in Panama, the Persian Gulf, Kosovo, Afghanistan, and Iraq. Israel used the Apache in its military conflicts in Lebanon and the Gaza Strip. British and Dutch Apaches have seen deployments in wars in Afghanistan and Iraq.