title: Revolutionary Road (film)

text: In mid-2007, the cast rehearsed for three-and-a-half weeks before principal photography and shot mostly in sequence and on location in Darien, Connecticut. Mendes wanted to create a claustrophobic dynamic on set, so he filmed the Wheeler home interiors in a real house. The property, including the neighbor's house, were very small but featured 1950s-style architecture. The homeowners gave permission for DreamWorks to dismantle and remodel the interior and exterior. Around 45 tradespeople were involved with the transformation, including carpenters, interior designers and landscapers. Production designer Kristi Zea and her team had five weeks to renovate the homes. "We wanted to keep the sense of isolation that was described in the book", Zea said. Debra Schutt served as a set decorator, and said, "We ruled out anything with bright colors and anything that hit you over the head with the period. The look is really quite plain".


title: Revolutionary Road (film)

text: Cinematographer Roger Deakins, who previously worked on Mendes' "Jarhead", shot with a combination of jib and handheld camera equipment. Deakins analyzed the use of every light fixture in the house; Schutt recalls "he’s quite specific about what he wants. For instance, for the night scene by the side of the road, he wanted streetlights that would give a rectangular, tapered light, and for an argument in the Wheelers’ front room, he wanted a ceiling fixture that would send light down and out in a fan shape with a hard edge". Period streetlights and automobiles were also fitted with specific light bulbs by the art department. Deakins found it difficult at times working in a small, shaded house with bulky camera equipment. Nevertheless, Winslet was impressed with his ability to "bounce light all over" using Arri Compact HMI lamps. To gradually illustrate the home's neglect and Wheelers' collapsing marriage, the crew removed props in the house and Deakins transitioned to handheld cameras, respectively. Mendes said, "I wanted a real rawness in Leo and Kate’s performances in the last half-hour of the movie, and when we reached that point, I told Roger I didn’t want to make any decisions [about shots]; I wanted it to be handheld, and I wanted to let the actors be explosive and unpredictable".


title: Revolutionary Road (film)

text: Recalling the on-set atmosphere, Michael Shannon said that he did not feel that there were any "stars", but "a group of people united by a passion for the material and wanting to honor the book". He said Winslet and DiCaprio could only make such a good performance as a couple because of their friendship since "Titanic". For Shannon, it was more important to prepare for the moment when he walked on the set than being concerned about the actors he was working with. In the fight scenes between DiCaprio and Winslet, DiCaprio said, "So much of what happens between Frank and April in this film is what's left unsaid. I actually found it a real joy to do those fight scenes because finally, these people were letting each other have it". Winslet described her working relationship with DiCaprio as "challenging" and "physically comfortable", but she also felt "pressure" working with him, and working with her then-husband Mendes. She added, "the on-set atmosphere was very fluid in that way in that we’d all share ideas [...] without treading on each other’s toes". DiCaprio found the filming process physically and emotionally exhausting, that he postponed his next film for two months.


title: Revolutionary Road (film)

text: Once filming was complete, Deakins had the film negative (Kodak Vision2 200T 5217 and 500T 5218) processed at DuArt Film and Video in New York, one of his favorite laboratories in the area. During post-production, handled by EFILM, Mendes deleted around 20 minutes of footage to keep it in the spirit of Yates' novel.


title: Revolutionary Road (film)

text: Thomas Newman composed the soundtrack for "Revolutionary Road"; it is his fourth film score collaboration with Mendes. Consisting of fifteen tracks, Newman uses a variety of piano, strings, metallic sound effects and basslines for a haunting minimalist sound. The music was recorded at Newman Scoring Stage in Los Angeles and the album was released on December 23, 2008.


title: Revolutionary Road (film)

text: "Revolutionary Road" premiered in Los Angeles on December 15, 2008, followed by a limited U.S. release on December 26, 2008, and then a wider release (1,058 theaters) on January 23, 2009. In most other countries, it was released between January 15–30, 2009. The film earned $22.9 million at the domestic box office and $56.6 million internationally for a moderate worldwide $76 million.


title: Revolutionary Road (film)

text: "Revolutionary Road" was released on DVD and Blu-ray on June 2, 2009. The Blu-ray edition includes an audio commentary by Mendes, 26-minutes of deleted scenes, and two documentaries on the development of the project.


title: Revolutionary Road (film)

text: "Revolutionary Road" received generally positive reviews from critics. On the review aggregator Rotten Tomatoes, the film has an approval rating of 67% based on 215 reviews, with an average score of 6.60/10. The website's critical consensus reads, "Brilliantly acted and emotionally powerful, "Revolutionary Road" is a handsome adaptation of Richard Yates' celebrated novel". On Metacritic, the film has a weighted average score of 69 out of 100, based on 38 critics, indicating "generally favorable reviews".


title: Revolutionary Road (film)

text: Rex Reed of "The New York Observer" gave the film a positive response; "a flawless, moment-to-moment autopsy of a marriage on the rocks and an indictment of the American Dream gone sour" and "a profound, intelligent and deeply heartfelt work that raises the bar of filmmaking to exhilarating". Peter Travers of "Rolling Stone" called the film "raw and riveting" and commented, "Directed with extraordinary skill by Sam Mendes, who warms the chill in the Yates-faithful script by Justin Haythe, the film is a tough road well worth traveling ... DiCaprio is in peak form, bringing layers of buried emotion to a defeated man. And the glorious Winslet defines what makes an actress great, blazing commitment to a character and the range to make every nuance felt". Writing for the "San Francisco Chronicle," Mick LaSalle said, "Finally, this is a movie that can and should be seen more than once. Watch it one time through her eyes. Watch it again through his eyes. It works both ways. It works in every way. This is a great American film".


title: Revolutionary Road (film)

text: Roger Ebert of the "Chicago Sun-Times" gave "Revolutionary Road" a maximum rating of four stars, commending the acting and screenplay and calling the film "so good it is devastating". Of DiCaprio and Winslet, he said, "they are so good, they stop being actors and become the people I grew up around". "Entertainment Weekly's" Owen Gleiberman graded the film B+ and commented, "The film is lavishly dark—some might say too dark—yet I'd suggest it has a different limitation: For all its shattering domestic discord, there's something remote and aestheticized about it. April brings a private well of conflict to her middle-class prison, but Winslet is so meticulous in her telegraphed despair that she intrigues us, moves us, yet never quite touches our unguarded nerves". Joe Neumaier of the "New York Daily News" said:


title: Revolutionary Road (film)

text: Some film critics gave a mixed response. David Ansen of "Newsweek" opined that it is "impeccably mounted—perhaps too much so. Mendes [...] has an innately theatrical style: everything pops off the screen a little bigger and bolder than life [...] Instead of losing myself in the story, I often felt on the outside looking in, appreciating the craftsmanship, but one step removed from the agony on display. "Revolutionary Road" is impressive, but it feels like a classic encased in amber". Kirk Honeycutt of "The Hollywood Reporter" described the film as a "didactic, emotionally overblown critique of the soulless suburbs", and thought it was a repeat of "American Beauty." He wrote, "Once more, the suburbs are well-upholstered nightmares and its denizens clueless—other than one estranged male. Everything is boldly indicated to the audience from arch acting styles to the wink-wink, nod-nod of its design. Indeed his actors play the subtext with such fury that the text virtually disappears. Subtlety is not one of Mendes' strong suits".


title: Revolutionary Road (film)

text: Writing for "Variety" magazine, Todd McCarthy thought the film was "faithful, intelligent, admirably acted, superbly shot". He added, "It also offers a near-perfect case study of the ways in which film is incapable of capturing certain crucial literary qualities, in this case the very things that elevate the book from being a merely insightful study of a deteriorating marriage into a remarkable one [...] Even when the dramatic temperature is cranked up too high, the picture's underpinnings seem only partly present, to the point where one suspects that what it's reaching for dramatically might be all but unattainable—perhaps approachable only by Pinter at his peak." McCarthy later changed his opinion, calling "Revolutionary Road" "problematic" and that it "has some issues that just won't go away". He concludes that "Revolutionary Road" suffers in comparison to Billy Wilder's "The Apartment" and Richard Quine's "Strangers When We Meet" because of its "narrow vision", even arguing that the television series "Mad Men" handles the issues of conformity, frustration, and hypocrisy "with more panache and precision".


title: BTR-80

text: The BTR-80 (, literally "armoured transporter") is an 8×8 wheeled amphibious armoured personnel carrier (APC) designed in the USSR. It was adopted in 1985 and replaced the previous vehicles, the BTR-60 and BTR-70, in the Soviet Army. It was first deployed during the Soviet–Afghan War.


title: BTR-80

text: The Soviets based the BTR-80 on the BTR-70 APC, which itself was based on the BTR-60. It has a single 260-hp V-8 turbocharged water-cooled diesel engine, an improvement over the twin gasoline engines installed in the BTR-60 and BTR-70 vehicles. The reconfigured rear portion of the hull accommodates the new, single engine. The Soviets removed the roof chamfers of the modified BTR-70, raised the rear, and squared off the rearward-sloping engine compartment.


title: BTR-80

text: Standard equipment includes TNPO vision blocks, TNP-B and TKN-3 optical devices for the driver and commander, an OU-3GA2M infrared search light, six 81 mm smoke grenade launchers 902V "Tucha", a radioset (R-173 or R-163-50U), an intercom, and hydrojets for amphibious propulsion.


title: BTR-80

text: The Soviets modified the truncated cone turret used on the BTR-70 for the BTR-80 by redesigning the mantlet. This allows the KPVT and coaxial PKT machine guns to be elevated to a maximum of 60 degrees. This high angle of fire is useful in engaging targets on steep slopes, in urban fighting, and for engaging low slow flying air targets.


title: BTR-80

text: The Soviets modified the design and positioning of the firing ports. The ports are now round rather than tear-shaped, and have ball mounts similar to those used on the BMP-1. The forward firing ports now sit in angled recesses, which allows infantry to fire to the front of the vehicle.


title: BTR-80

text: The redesigned side doors are split horizontally. The upper portion opens forward. This gives dismounting troops some protection against small arms fire from the front of the vehicle. The lower portion opens down, forming a step. Six smoke grenade projectors are mounted on the rear of the turret.


title: BTR-80

text: In 1984, the Soviets began production of a diesel-engined variant of the BTR-70, which they called the BTR-80. The Soviets have retrofitted some BTR-70s with several of the improvements incorporated into the BTR-80, including the high-angle-of-fire turret. The twin doors are designed to allow the infantry to disembark while the vehicle is in motion, and allow the infantry inside to exit from one side if the other is receiving fire.


title: BTR-80

text: The 30mm Cannon variants are effective against most targets apart from main battle tanks, against which they can still cause significant damage to optics, weapons, and important systems. The main gun is not stabilized, so accurate fire on the move is limited to low speeds. The turret's rotation mechanism is manually operated.


title: BTR-80

text: The gunner sits in a roof-mounted chair located above the flat floor behind the driver/commander and two passengers, and before the passenger bench. The gunner's station is basic, but uncharacteristically spacious for a Soviet armored vehicle. The gunner is equipped with both a day time optical sight and an infrared night sight.


title: BTR-80

text: The BTR-80 and BTR-82A took part in the 2022 Russian invasion of Ukraine. As of 13 November 2022, Russia was visually confirmed by Oryx to have lost 340 BTR-82As (213 destroyed, 6 damaged, 13 abandoned and 108 captured) and 132 BTR-80s (including 68 destroyed, 1 damaged, 4 abandoned and 59 captured). Another 44 BTR-80/BTR-82As were also destroyed or abandoned, but their models cannot be determined.


title: BTR-80

text: Nevertheless, the recognized Ukrainian involvement in the armament (ZTM-2 cannon) and the strong visual resemblance to BTR-3 hints us to state that it is (at least a derivative of) a BTR-3.


title: Barack Obama Sr.

text: Barack Hussein Obama Sr. (; 18 June 1934 – 24 November 1982) was a Kenyan senior governmental economist and the father of Barack Obama, the 44th president of the United States. He is a central figure of his son's memoir, "Dreams from My Father" (1995). Obama married in 1954 and had two children with his first wife, Kezia. He was selected for a special program to attend college in the United States and studied at the University of Hawaii where he met Stanley Ann Dunham, whom he married in 1961 following the conception of his son, Barack II. Dunham divorced Obama three years later. The elder Obama later went to Harvard University for graduate school, where he earned an M.A. in economics, and returned to Kenya in 1964. He saw his son Barack once more, when his son was about 10.


title: Barack Obama Sr.

text: In late 1964, Obama Sr. married Ruth Beatrice Baker, a Jewish-American woman he had met in Massachusetts. They had two sons together before separating in 1971 and divorcing in 1973. Obama first worked for an oil company, before beginning work as an economist with the Kenyan Ministry of Transport. He gained a promotion to senior economic analyst in the Ministry of Finance. He was among a cadre of young Kenyan men who had been educated in the West in a program supported by Tom Mboya. Obama Sr. had conflicts with Kenyan president Jomo Kenyatta, which adversely affected his career. He was fired and blacklisted in Kenya, finding it nearly impossible to get a job. Obama Sr. was involved in three serious car accidents during his final years; he died as a result of the last one in 1982.


title: Barack Obama Sr.

text: Barack Obama Sr. was born in 1934 in Rachuonyo District on the shores of Lake Victoria just outside Kendu Bay, British Kenya, at the time a colony and protectorate of the British Empire. He was raised in the village of Nyang'oma Kogelo, Siaya District, Nyanza Province. His family are members of the Luo ethnic group.


title: Barack Obama Sr.

text: His father was Onyango (later Hussein) Obama (c. 1895–1979), and his mother was Habiba Akumu Nyanjango (c. 1918–2006) of Karabondi, Kenya, Onyango's second wife. They had two daughters and a son together, Barack Obama (Sr.). After Akumu separated from Onyango and left the family in 1945, her three children were raised by Onyango's third wife, Sarah Ogwel of Kogelo.


title: Barack Obama Sr.

text: As a young man, Onyango Obama traveled widely, enlisting in the British colonial forces and visiting Europe, India, and Zanzibar. There, he converted from Roman Catholicism to Islam and took the name Hussein. He became a cook for missionaries and a local herbalist in Nairobi.


title: Barack Obama Sr.

text: "The Times" of London, relying on statements by Obama's third wife, Sarah Onyango Obama, 87, reported that in 1949, after becoming more politically active, Obama was jailed by the British for two years for suspicion of supplying military information to the Kenyan independence movement, owing to "His job as a cook to a British army officer." According to Sarah, Obama was subjected to beatings and abuse that left him with physical scars and a loathing of the British. David Maraniss' 2012 biography of President Barack Obama states that his grandfather Obama did not participate in the insurrections; nor was he ever imprisoned by the British during the uprising. He continued to be trusted by white Kenyans.


title: Barack Obama Sr.

text: When Obama (Sr.) was about six years old and attending a Christian missionary school, he converted from Islam to Anglicanism when strongly encouraged by the staff. He changed his name from "Baraka" to "Barack". Obama (Sr.) later became an atheist, believing that religion was mere superstition.


title: Barack Obama Sr.

text: While still living near Kendu Bay, Obama (Sr.) attended Gendia Primary School. After his family moved to Siaya District, he transferred to Ng'iya Intermediate School. From 1950 to 1953, he studied at Maseno National School, an exclusive Anglican boarding school in Maseno. The head teacher, B.L. Bowers, described Obama in his records as "very keen, steady, trustworthy and friendly. Concentrates, reliable and out-going."


title: Barack Obama Sr.

text: In 1954 at age 20, Obama (Sr.) married Kezia Aoko in a tribal ceremony in Kenya. They had two children, Malik (a.k.a. Roy) and Auma, during the early years of their marriage.


title: Barack Obama Sr.

text: Obama (Sr.) married two other women. In 1960 while studying at the University of Hawai'i on a special government-sponsored program, he met and dated Stanley Ann Dunham, an American woman. She became pregnant and they were married on the Hawaiian island of Maui on 2 February 1961, despite parental opposition from both families. Their son, future US president Barack Obama, was born in August 1961. They later separated and divorced on 20 March 1964, whereupon Ann Dunham was granted sole custody of their son.


title: Barack Obama Sr.

text: After his 1962 graduation from the University of Hawaii, Obama Sr. went to Harvard University for a doctoral program in economics, but left the university after completing his master's degree. At Harvard he started dating Ruth Beatrice Baker, a Jewish-American woman. She went with him to Kenya in 1964. They married late that year and had two sons, Mark and David. The couple separated in 1971 and divorced in 1973.


title: Barack Obama Sr.

text: In Kenya, Obama Sr. reconnected with his first wife Kezia. She had two sons after his return: Abo (b. 1968) and Bernard (b. 1970), believed to be his children.


title: Barack Obama Sr.

text: Barack Obama, in his memoir "Dreams from My Father" (1995), said that his father's family had questioned whether Abo and Bernard are Barack Sr.'s biological sons.


title: Barack Obama Sr.

text: The senior Obama fathered his last son George in 1982 with Jael Otinyo. George was about six months old when Obama Sr. was killed in a car accident. George was raised by his mother, who later remarried; his stepfather cared for him as well.


title: Barack Obama Sr.

text: In 1959, the Kenyan Department of Education published Obama's monograph, entitled "Otieno jarieko. Kitabu mar ariyo. 2: Yore mabeyo mag puro puothe." (English: "Otieno, the wise man. Book 2: Wise ways of farming.")


title: Barack Obama Sr.

text: Due to his accomplishments, in 1959 Obama received a scholarship in economics through a program organized by the nationalist leader Tom Mboya. The program offered education in the West to outstanding Kenyan students. Initial financial supporters of the program included Harry Belafonte, Sidney Poitier, Jackie Robinson, and Elizabeth Mooney Kirk, a literacy advocate who provided most of the financial support for Obama's early years in the United States. Kirk and her literacy associate Helen M. Roberts of Palo Alto raised the money necessary for Obama to travel to the US.


title: Barack Obama Sr.

text: When Obama left for the United States, he left behind his young wife, Kezia, and their baby son Malik. Kezia was pregnant, and their daughter Auma was born while her father was in Hawaii. At Obama's request, Helen Roberts committed to watching over and financially supporting the family that he had left behind, for as long as she remained in Nairobi.


title: Barack Obama Sr.

text: In 1959, Obama enrolled at the University of Hawaii at Manoa in Honolulu as the university's first African foreign student. He initially lived across the street from the university at the Charles H. Atherton branch of the YMCA at 1810 University Avenue; public records from 1961 indicate he later had a residence two miles southeast of the university at 625 11th Avenue in the Kaimuki neighborhood.


title: Barack Obama Sr.

text: In 1960, Obama met Stanley Ann Dunham in a basic Russian language course at the University of Hawaii and they started dating. After becoming pregnant, Dunham dropped out of the University of Hawaii after the fall 1960 semester, while Obama continued his education. Obama married Dunham in Wailuku on the Hawaiian island of Maui on 2 February 1961. He eventually told Dunham about his previous marriage in Kenya, but said he was divorced—which she found out years later was not true.


title: Barack Obama Sr.

text: Their son Barack Obama II, was born in Honolulu on 4 August 1961 at the former Kapiolani Maternity and Gynecological Hospital (succeeded by the Kapi'olani Medical Center for Women and Children). His birth was announced in "The Honolulu Advertiser" and the "Honolulu Star-Bulletin," with his parents' address listed as 6085 Kalanianaole Highway in the Kuliouou neighborhood of Honolulu, seven miles east of the university—the rented home of Dunham's parents, Stanley and Madelyn Dunham. Soon after her son's birth, Dunham took the infant with her to Seattle, Washington, where she took classes at the University of Washington from September 1961 to June 1962.


title: Barack Obama Sr.

text: Obama Sr. continued his education at the University of Hawaii and in 1961–1962 lived one mile east of the university in the St. Louis Heights neighborhood. He graduated from the University of Hawaii after three years with a B.A. in economics and was elected to Phi Beta Kappa. He left Hawaii in June 1962.


title: Barack Obama Sr.

text: In September 1962, after a tour of mainland U.S. universities, Obama Sr. traveled to Cambridge, Massachusetts, where he began a graduate fellowship in economics at Harvard University. He rented an apartment in a rooming house near Central Square in Cambridge. Meanwhile, Dunham and their son returned to Honolulu in the latter half of 1962, and she resumed her undergraduate education in January 1963 in the spring semester at the University of Hawaii. In January 1964, Dunham filed for divorce in Honolulu; the divorce was not contested by Obama. In 1965, Dunham married Lolo Soetoro, a Javanese surveyor whom she had met at the East-West Center.


title: Barack Obama Sr.

text: Obama was forced to leave his PhD program at Harvard University in May 1964 (and received an M.A. in economics from Harvard in 1965). In June 1964, Obama met and began dating a 27-year-old Jewish-American elementary school teacher named Ruth Beatrice Baker, the daughter of prosperous Lithuanian immigrants to the United States.


title: Barack Obama Sr.

text: Obama returned to Kenya in 1964 after graduating from Harvard. Baker followed him, and they married 24 December 1964. They had two sons together, Mark Okoth Obama in 1965 and David Opiyo Obama in 1968. Baker and Obama separated in 1971, and divorced in 1973. Baker subsequently married a Tanzanian man named Ndesandjo and took his surname, as did her sons Mark and David. Mark said in 2009 that Obama had been abusive to him, his late brother David, and their mother.


title: Barack Obama Sr.

text: Obama first worked as a government economist for an oil company in Kenya. In 1965, Obama published a paper entitled "Problems Facing Our Socialism" in the "East Africa Journal," harshly criticizing the blueprint for national planning, "African Socialism and Its Applicability to Planning in Kenya", developed by Tom Mboya's Ministry of Economic Planning and Development. Obama served as an economist in the Kenyan Ministry of Transport. Later he was promoted to senior economist in the Kenyan Ministry of Finance.


title: Barack Obama Sr.

text: In 1970, Obama was in a serious automobile accident, and was hospitalized for nearly a year. In December 1971, he traveled to Hawaii for a month. There he visited with his ex-wife Ann Dunham and American son Barack II, who was nearly 10. The visit was the last time the boy saw his father. During his trip, Obama took his son to his first jazz concert, a performance by the pianist Dave Brubeck.


title: Barack Obama Sr.

text: According to Barack II's memoir, Obama's continuing conflict with Kenyan president Jomo Kenyatta destroyed his career. He came under suspicion after Tom Mboya was assassinated in 1969, as he had been a protege of the ruler. Kenyatta fired Obama, who was blacklisted in Kenya and found it impossible to get work. By the time Obama visited his son in Hawaii in 1971, he had a bad leg from the 1970 accident.


title: Barack Obama Sr.

text: Obama later lost both legs in a second serious automobile accident, and subsequently lost his job. His life deteriorated as he struggled with poverty and drinking. During his final decade, he never recovered his former social or economic standing. His friend Philip Ochieng, a journalist of the Kenya newspaper "Daily Nation," has described Obama's difficult personality and drinking problems. In 1982, Obama had a relationship with Jael Otinyo and with her fathered his last son, named George Obama. Six months after George's birth, Obama died in a car crash in Nairobi. He was interred in his native village of Nyang'oma Kogelo, Siaya District. His funeral was attended by ministers Robert Ouko, Peter Oloo-Aringo, and other prominent political figures.


title: Beetle

text: Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 described species, is the largest of all orders, constituting almost 40% of described insects and 25% of all known animal life-forms; new species are discovered frequently, with estimates suggesting that there are between 0.9 and 2.1 million total species. Found in almost every habitat except the sea and the polar regions, they interact with their ecosystems in several ways: beetles often feed on plants and fungi, break down animal and plant debris, and eat other invertebrates. Some species are serious agricultural pests, such as the Colorado potato beetle, while others such as Coccinellidae (ladybirds or ladybugs) eat aphids, scale insects, thrips, and other plant-sucking insects that damage crops.


title: Beetle

text: Beetles typically have a particularly hard exoskeleton including the elytra, though some such as the rove beetles have very short elytra while blister beetles have softer elytra. The general anatomy of a beetle is quite uniform and typical of insects, although there are several examples of novelty, such as adaptations in water beetles which trap air bubbles under the elytra for use while diving. Beetles are endopterygotes, which means that they undergo complete metamorphosis, with a series of conspicuous and relatively abrupt changes in body structure between hatching and becoming adult after a relatively immobile pupal stage. Some, such as stag beetles, have a marked sexual dimorphism, the males possessing enormously enlarged mandibles which they use to fight other males. Many beetles are aposematic, with bright colors and patterns warning of their toxicity, while others are harmless Batesian mimics of such insects. Many beetles, including those that live in sandy places, have effective camouflage.


title: Beetle

text: Beetles are prominent in human culture, from the sacred scarabs of ancient Egypt to beetlewing art and use as pets or fighting insects for entertainment and gambling. Many beetle groups are brightly and attractively colored making them objects of collection and decorative displays. Over 300 species are used as food, mostly as larvae; species widely consumed include mealworms and rhinoceros beetle larvae. However, the major impact of beetles on human life is as agricultural, forestry, and horticultural pests. Serious pests include the boll weevil of cotton, the Colorado potato beetle, the coconut hispine beetle, and the mountain pine beetle. Most beetles, however, do not cause economic damage and many, such as the lady beetles and dung beetles are beneficial by helping to control insect pests.


title: Beetle

text: The name of the taxonomic order, Coleoptera, comes from the Greek "koleopteros" (κολεόπτερος), given to the group by Aristotle for their elytra, hardened shield-like forewings, from "koleos", sheath, and "pteron", wing. The English name beetle comes from the Old English word "bitela", little biter, related to "bītan" (to bite), leading to Middle English "betylle". Another Old English name for beetle is "ċeafor", chafer, used in names such as cockchafer, from the Proto-Germanic *"kebrô" ("beetle"; compare German "Käfer", Dutch "kever").


title: Beetle

text: Beetles are by far the largest order of insects: the roughly 400,000 species make up about 40% of all insect species so far described, and about 25% of all animals. A 2015 study provided four independent estimates of the total number of beetle species, giving a mean estimate of some 1.5 million with a "surprisingly narrow range" spanning all four estimates from a minimum of 0.9 to a maximum of 2.1 million beetle species. The four estimates made use of host-specificity relationships (1.5 to 1.9 million), ratios with other taxa (0.9 to 1.2 million), plant:beetle ratios (1.2 to 1.3), and extrapolations based on body size by year of description (1.7 to 2.1 million).


title: Beetle

text: Beetles are found in nearly all habitats, including freshwater and coastal habitats, wherever vegetative foliage is found, from trees and their bark to flowers, leaves, and underground near roots - even inside plants in galls, in every plant tissue, including dead or decaying ones. Tropical forest canopies have a large and diverse fauna of beetles, including Carabidae, Chrysomelidae, and Scarabaeidae.


title: Beetle

text: The heaviest beetle, indeed the heaviest insect stage, is the larva of the goliath beetle, "Goliathus goliatus", which can attain a mass of at least and a length of . Adult male goliath beetles are the heaviest beetle in its adult stage, weighing and measuring up to . Adult elephant beetles, "Megasoma elephas" and "Megasoma actaeon" often reach and .


title: Beetle

text: The longest beetle is the Hercules beetle "Dynastes hercules", with a maximum overall length of at least 16.7 cm (6.6 in) including the very long pronotal horn. The smallest recorded beetle and the smallest free-living insect (), is the featherwing beetle "Scydosella musawasensis" which may measure as little as 325 μm in length.


title: Beetle

text: The oldest known beetle is "Coleopsis", from the earliest Permian (Asselian) of Germany, around 295 million years ago. Early beetles from the Permian, which are collectively grouped into the "Protocoleoptera" are thought to have been xylophagous (wood eating) and wood boring. Fossils from this time have been found in Siberia and Europe, for instance in the red slate fossil beds of Niedermoschel near Mainz, Germany. Further fossils have been found in Obora, Czech Republic and Tshekarda in the Ural mountains, Russia. However, there are only a few fossils from North America before the middle Permian, although both Asia and North America had been united to Euramerica. The first discoveries from North America made in the Wellington Formation of Oklahoma were published in 2005 and 2008. The earliest members of modern beetle lineages appeared during the Late Permian. In the Permian–Triassic extinction event at the end of the Permian, most "protocoleopteran" lineages became extinct. Beetle diversity did not recover to pre-extinction levels until the Middle Triassic.


title: Beetle

text: During the Jurassic (), there was a dramatic increase in the diversity of beetle families, including the development and growth of carnivorous and herbivorous species. The Chrysomeloidea diversified around the same time, feeding on a wide array of plant hosts from cycads and conifers to angiosperms. Close to the Upper Jurassic, the Cupedidae decreased, but the diversity of the early plant-eating species increased. Most recent plant-eating beetles feed on flowering plants or angiosperms, whose success contributed to a doubling of plant-eating species during the Middle Jurassic. However, the increase of the number of beetle families during the Cretaceous does not correlate with the increase of the number of angiosperm species. Around the same time, numerous primitive weevils (e.g. Curculionoidea) and click beetles (e.g. Elateroidea) appeared. The first jewel beetles (e.g. Buprestidae) are present, but they remained rare until the Cretaceous. The first scarab beetles were not coprophagous but presumably fed on rotting wood with the help of fungus; they are an early example of a mutualistic relationship.


title: Beetle

text: There are more than 150 important fossil sites from the Jurassic, the majority in Eastern Europe and North Asia. Outstanding sites include Solnhofen in Upper Bavaria, Germany, Karatau in South Kazakhstan, the Yixian formation in Liaoning, North China, as well as the Jiulongshan formation and further fossil sites in Mongolia. In North America there are only a few sites with fossil records of insects from the Jurassic, namely the shell limestone deposits in the Hartford basin, the Deerfield basin and the Newark basin.


title: Beetle

text: The Cretaceous saw the fragmenting of the southern landmass, with the opening of the southern Atlantic Ocean and the isolation of New Zealand, while South America, Antarctica, and Australia grew more distant. The diversity of Cupedidae and Archostemata decreased considerably. Predatory ground beetles (Carabidae) and rove beetles (Staphylinidae) began to distribute into different patterns; the Carabidae predominantly occurred in the warm regions, while the Staphylinidae and click beetles (Elateridae) preferred temperate climates. Likewise, predatory species of Cleroidea and Cucujoidea hunted their prey under the bark of trees together with the jewel beetles (Buprestidae). The diversity of jewel beetles increased rapidly, as they were the primary consumers of wood, while longhorn beetles (Cerambycidae) were rather rare: their diversity increased only towards the end of the Upper Cretaceous. The first coprophagous beetles are from the Upper Cretaceous and may have lived on the excrement of herbivorous dinosaurs. The first species where both larvae and adults are adapted to an aquatic lifestyle are found. Whirligig beetles (Gyrinidae) were moderately diverse, although other early beetles (e.g. Dytiscidae) were less, with the most widespread being the species of Coptoclavidae, which preyed on aquatic fly larvae.


title: Beetle

text: A 2020 review of the palaeoecological interpretations of fossil beetles from Cretaceous ambers has suggested that saproxylicity was the most common feeding strategy, with fungivorous species in particular appearing to dominate.


title: Beetle

text: Many fossil sites worldwide contain beetles from the Cretaceous. Most are in Europe and Asia and belong to the temperate climate zone during the Cretaceous. Lower Cretaceous sites include the Crato fossil beds in the Araripe basin in the Ceará, North Brazil, as well as overlying Santana formation; the latter was near the equator at that time. In Spain, important sites are near Montsec and Las Hoyas. In Australia, the Koonwarra fossil beds of the Korumburra group, South Gippsland, Victoria, are noteworthy. Major sites from the Upper Cretaceous include Kzyl-Dzhar in South Kazakhstan and Arkagala in Russia.


title: Beetle

text: Beetle fossils are abundant in the Cenozoic; by the Quaternary (up to 1.6 mya), fossil species are identical to living ones, while from the Late Miocene (5.7 mya) the fossils are still so close to modern forms that they are most likely the ancestors of living species. The large oscillations in climate during the Quaternary caused beetles to change their geographic distributions so much that current location gives little clue to the biogeographical history of a species. It is evident that geographic isolation of populations must often have been broken as insects moved under the influence of changing climate, causing mixing of gene pools, rapid evolution, and extinctions, especially in middle latitudes.


title: Beetle

text: The very large number of beetle species poses special problems for classification. Some families contain tens of thousands of species, and need to be divided into subfamilies and tribes. This immense number led the evolutionary biologist J. B. S. Haldane to quip, when some theologians asked him what could be inferred about the mind of the Christian God from the works of His Creation, "An inordinate fondness for beetles". Polyphaga is the largest suborder, containing more than 300,000 described species in more than 170 families, including rove beetles (Staphylinidae), scarab beetles (Scarabaeidae), blister beetles (Meloidae), stag beetles (Lucanidae) and true weevils (Curculionidae). These polyphagan beetle groups can be identified by the presence of cervical sclerites (hardened parts of the head used as points of attachment for muscles) absent in the other suborders.


title: Beetle

text: Adephaga contains about 10 families of largely predatory beetles, includes ground beetles (Carabidae), water beetles (Dytiscidae) and whirligig beetles (Gyrinidae). In these insects, the testes are tubular and the first abdominal sternum (a plate of the exoskeleton) is divided by the hind coxae (the basal joints of the beetle's legs).


title: Beetle

text: Archostemata contains four families of mainly wood-eating beetles, including reticulated beetles (Cupedidae) and the telephone-pole beetle.


title: Beetle

text: The Archostemata have an exposed plate called the metatrochantin in front of the basal segment or coxa of the hind leg. Myxophaga contains about 65 described species in four families, mostly very small, including Hydroscaphidae and the genus "Sphaerius". The myxophagan beetles are small and mostly alga-feeders. Their mouthparts are characteristic in lacking galeae and having a mobile tooth on their left mandible.


title: Beetle

text: The consistency of beetle morphology, in particular their possession of elytra, has long suggested that Coleoptera is monophyletic, though there have been doubts about the arrangement of the suborders, namely the Adephaga, Archostemata, Myxophaga and Polyphaga within that clade. The twisted-wing parasites, Strepsiptera, are thought to be a sister group to the beetles, having split from them in the Early Permian.


title: Beetle

text: Molecular phylogenetic analysis confirms that the Coleoptera are monophyletic. Duane McKenna et al. (2015) used eight nuclear genes for 367 species from 172 of 183 Coleopteran families. They split the Adephaga into 2 clades, Hydradephaga and Geadephaga, broke up the Cucujoidea into 3 clades, and placed the Lymexyloidea within the Tenebrionoidea. The Polyphaga appear to date from the Triassic. Most extant beetle families appear to have arisen in the Cretaceous. The cladogram is based on McKenna (2015). The number of species in each group (mainly superfamilies) is shown in parentheses, and boldface if over 10,000. English common names are given where possible. Dates of origin of major groups are shown in italics in millions of years ago (mya).


title: Beetle

text: Beetles are generally characterized by a particularly hard exoskeleton and hard forewings (elytra) not usable for flying. Almost all beetles have mandibles that move in a horizontal plane. The mouthparts are rarely suctorial, though they are sometimes reduced; the maxillae always bear palps. The antennae usually have 11 or fewer segments, except in some groups like the Cerambycidae (longhorn beetles) and the Rhipiceridae (cicada parasite beetles). The coxae of the legs are usually located recessed within a coxal cavity. The genitalic structures are telescoped into the last abdominal segment in all extant beetles. Beetle larvae can often be confused with those of other endopterygote groups. The beetle's exoskeleton is made up of numerous plates, called sclerites, separated by thin sutures. This design provides armored defenses while maintaining flexibility. The general anatomy of a beetle is quite uniform, although specific organs and appendages vary greatly in appearance and function between the many families in the order. Like all insects, beetles' bodies are divided into three sections: the head, the thorax, and the abdomen. Because there are so many species, identification is quite difficult, and relies on attributes including the shape of the antennae, the tarsal formulae and shapes of these small segments on the legs, the mouthparts, and the ventral plates (sterna, pleura, coxae). In many species accurate identification can only be made by examination of the unique male genitalic structures.


title: Beetle

text: The head, having mouthparts projecting forward or sometimes downturned, is usually heavily sclerotized and is sometimes very large. The eyes are compound and may display remarkable adaptability, as in the case of the aquatic whirligig beetles (Gyrinidae), where they are split to allow a view both above and below the waterline. A few Longhorn beetles (Cerambycidae) and weevils as well as some fireflies (Rhagophthalmidae) have divided eyes, while many have eyes that are notched, and a few have ocelli, small, simple eyes usually farther back on the head (on the vertex); these are more common in larvae than in adults. The anatomical organization of the compound eyes may be modified and depends on whether a species is primarily crepuscular, or diurnally or nocturnally active. Ocelli are found in the adult carpet beetle (Dermestidae), some rove beetles (Omaliinae), and the Derodontidae.


title: Beetle

text: Beetle antennae are primarily organs of sensory perception and can detect motion, odor and chemical substances, but may also be used to physically feel a beetle's environment. Beetle families may use antennae in different ways. For example, when moving quickly, tiger beetles may not be able to see very well and instead hold their antennae rigidly in front of them in order to avoid obstacles.


title: Beetle

text: Certain Cerambycidae use antennae to balance, and blister beetles may use them for grasping. Some aquatic beetle species may use antennae for gathering air and passing it under the body whilst submerged. Equally, some families use antennae during mating, and a few species use them for defense. In the cerambycid "Onychocerus albitarsis", the antennae have venom injecting structures used in defense, which is unique among arthropods. Antennae vary greatly in form, sometimes between the sexes, but are often similar within any given family. Antennae may be , , , , (either on one side or both, bipectinate), or . The physical variation of antennae is important for the identification of many beetle groups. The Curculionidae have elbowed or geniculate antennae. Feather like flabellate antennae are a restricted form found in the Rhipiceridae and a few other families. The Silphidae have a capitate antennae with a spherical head at the tip. The Scarabaeidae typically have lamellate antennae with the terminal segments extended into long flat structures stacked together. The Carabidae typically have thread-like antennae. The antennae arises between the eye and the mandibles and in the Tenebrionidae, the antennae rise in front of a notch that breaks the usually circular outline of the compound eye. They are segmented and usually consist of 11 parts, the first part is called the scape and the second part is the pedicel. The other segments are jointly called the flagellum.


title: Beetle

text: Beetles have mouthparts like those of grasshoppers. The mandibles appear as large pincers on the front of some beetles. The mandibles are a pair of hard, often tooth-like structures that move horizontally to grasp, crush, or cut food or enemies (see defence, below). Two pairs of finger-like appendages, the maxillary and labial palpi, are found around the mouth in most beetles, serving to move food into the mouth. In many species, the mandibles are sexually dimorphic, with those of the males enlarged enormously compared with those of females of the same species.


title: Beetle

text: The thorax is segmented into the two discernible parts, the pro- and pterothorax. The pterothorax is the fused meso- and metathorax, which are commonly separated in other insect species, although flexibly articulate from the prothorax. When viewed from below, the thorax is that part from which all three pairs of legs and both pairs of wings arise. The abdomen is everything posterior to the thorax. When viewed from above, most beetles appear to have three clear sections, but this is deceptive: on the beetle's upper surface, the middle section is a hard plate called the pronotum, which is only the front part of the thorax; the back part of the thorax is concealed by the beetle's wings. This further segmentation is usually best seen on the abdomen.


title: Beetle

text: The multisegmented legs end in two to five small segments called tarsi. Like many other insect orders, beetles have claws, usually one pair, on the end of the last tarsal segment of each leg. While most beetles use their legs for walking, legs have been variously adapted for other uses. Aquatic beetles including the Dytiscidae (diving beetles), Haliplidae, and many species of Hydrophilidae, the legs, often the last pair, are modified for swimming, typically with rows of long hairs. Male diving beetles have suctorial cups on their forelegs that they use to grasp females. Other beetles have fossorial legs widened and often spined for digging. Species with such adaptations are found among the scarabs, ground beetles, and clown beetles (Histeridae). The hind legs of some beetles, such as flea beetles (within Chrysomelidae) and flea weevils (within Curculionidae), have enlarged femurs that help them leap.


title: Beetle

text: The forewings of beetles are not used for flight, but form elytra which cover the hind part of the body and protect the hindwings. The elytra are usually hard shell-like structures which must be raised to allow the hindwings to move for flight. However, in the soldier beetles (Cantharidae), the elytra are soft, earning this family the name of leatherwings. Other soft wing beetles include the net-winged beetle "Calopteron discrepans", which has brittle wings that rupture easily in order to release chemicals for defense.


title: Beetle

text: Beetles' flight wings are crossed with veins and are folded after landing, often along these veins, and stored below the elytra. A fold ("jugum") of the membrane at the base of each wing is characteristic. Some beetles have lost the ability to fly. These include some ground beetles (Carabidae) and some true weevils (Curculionidae), as well as desert- and cave-dwelling species of other families. Many have the two elytra fused together, forming a solid shield over the abdomen. In a few families, both the ability to fly and the elytra have been lost, as in the glow-worms (Phengodidae), where the females resemble larvae throughout their lives. The presence of elytra and wings does not always indicate that the beetle will fly. For example, the tansy beetle walks between habitats despite being physically capable of flight.


title: Beetle

text: The abdomen is the section behind the metathorax, made up of a series of rings, each with a hole for breathing and respiration, called a spiracle, composing three different segmented sclerites: the tergum, pleura, and the sternum. The tergum in almost all species is membranous, or usually soft and concealed by the wings and elytra when not in flight. The pleura are usually small or hidden in some species, with each pleuron having a single spiracle. The sternum is the most widely visible part of the abdomen, being a more or less sclerotized segment. The abdomen itself does not have any appendages, but some (for example, Mordellidae) have articulating sternal lobes.


title: Beetle

text: The digestive system of beetles is primarily adapted for a herbivorous diet. Digestion takes place mostly in the anterior midgut, although in predatory groups like the Carabidae, most digestion occurs in the crop by means of midgut enzymes. In the Elateridae, the larvae are liquid feeders that extraorally digest their food by secreting enzymes. The alimentary canal basically consists of a short, narrow pharynx, a widened expansion, the crop, and a poorly developed gizzard. This is followed by the midgut, that varies in dimensions between species, with a large amount of cecum, and the hindgut, with varying lengths. There are typically four to six Malpighian tubules.


title: Beetle

text: The nervous system in beetles contains all the types found in insects, varying between different species, from three thoracic and seven or eight abdominal ganglia which can be distinguished to that in which all the thoracic and abdominal ganglia are fused to form a composite structure.


title: Beetle

text: Like most insects, beetles inhale air, for the oxygen it contains, and exhale carbon dioxide, via a tracheal system. Air enters the body through spiracles, and circulates within the haemocoel in a system of tracheae and tracheoles, through whose walls the gases can diffuse.


title: Beetle

text: Diving beetles, such as the Dytiscidae, carry a bubble of air with them when they dive. Such a bubble may be contained under the elytra or against the body by specialized hydrophobic hairs. The bubble covers at least some of the spiracles, permitting air to enter the tracheae. The function of the bubble is not only to contain a store of air but to act as a physical gill. The air that it traps is in contact with oxygenated water, so as the animal's consumption depletes the oxygen in the bubble, more oxygen can diffuse in to replenish it. Carbon dioxide is more soluble in water than either oxygen or nitrogen, so it readily diffuses out faster than in. Nitrogen is the most plentiful gas in the bubble, and the least soluble, so it constitutes a relatively static component of the bubble and acts as a stable medium for respiratory gases to accumulate in and pass through. Occasional visits to the surface are sufficient for the beetle to re-establish the constitution of the bubble.


title: Beetle

text: Like other insects, beetles have open circulatory systems, based on hemolymph rather than blood. As in other insects, a segmented tube-like heart is attached to the dorsal wall of the hemocoel. It has paired inlets or "ostia" at intervals down its length, and circulates the hemolymph from the main cavity of the haemocoel and out through the anterior cavity in the head.


title: Beetle

text: Different glands are specialized for different pheromones to attract mates. Pheromones from species of Rutelinae are produced from epithelial cells lining the inner surface of the apical abdominal segments; amino acid-based pheromones of Melolonthinae are produced from eversible glands on the abdominal apex. Other species produce different types of pheromones. Dermestids produce esters, and species of Elateridae produce fatty acid-derived aldehydes and acetates. To attract a mate, fireflies (Lampyridae) use modified fat body cells with transparent surfaces backed with reflective uric acid crystals to produce light by bioluminescence. Light production is highly efficient, by oxidation of luciferin catalyzed by enzymes (luciferases) in the presence of adenosine triphosphate (ATP) and oxygen, producing oxyluciferin, carbon dioxide, and light.


title: Beetle

text: Tympanal organs or hearing organs consist of a membrane (tympanum) stretched across a frame backed by an air sac and associated sensory neurons, are found in two families. Several species of the genus "Cicindela" (Carabidae) have hearing organs on the dorsal surfaces of their first abdominal segments beneath the wings; two tribes in the Dynastinae (within the Scarabaeidae) have hearing organs just beneath their pronotal shields or neck membranes. Both families are sensitive to ultrasonic frequencies, with strong evidence indicating they function to detect the presence of bats by their ultrasonic echolocation.


title: Beetle

text: Beetles are members of the superorder Endopterygota, and accordingly most of them undergo complete metamorphosis. The typical form of metamorphosis in beetles passes through four main stages: the egg, the larva, the pupa, and the imago or adult. The larvae are commonly called grubs and the pupa sometimes is called the chrysalis. In some species, the pupa may be enclosed in a cocoon constructed by the larva towards the end of its final instar. Some beetles, such as typical members of the families Meloidae and Rhipiphoridae, go further, undergoing hypermetamorphosis in which the first instar takes the form of a triungulin.


title: Beetle

text: Some beetles have intricate mating behaviour. Pheromone communication is often important in locating a mate.


title: Beetle

text: Different species use different pheromones. Scarab beetles such as the Rutelinae use pheromones derived from fatty acid synthesis, while other scarabs such as the Melolonthinae use amino acids and terpenoids. Another way beetles find mates is seen in the fireflies (Lampyridae) which are bioluminescent, with abdominal light-producing organs. The males and females engage in a complex dialog before mating; each species has a unique combination of flight patterns, duration, composition, and intensity of the light produced.


title: Beetle

text: Before mating, males and females may stridulate, or vibrate the objects they are on. In the Meloidae, the male climbs onto the dorsum of the female and strokes his antennae on her head, palps, and antennae. In "Eupompha", the male draws his antennae along his longitudinal vertex. They may not mate at all if they do not perform the precopulatory ritual. This mating behavior may be different amongst dispersed populations of the same species. For example, the mating of a Russian population of tansy beetle ("Chysolina graminis") is preceded by an elaborate ritual involving the male tapping the female's eyes, pronotum and antennae with its antennae, which is not evident in the population of this species in the United Kingdom.


title: Beetle

text: Competition can play a part in the mating rituals of species such as burying beetles ("Nicrophorus"), the insects fighting to determine which can mate. Many male beetles are territorial and fiercely defend their territories from intruding males. In such species, the male often has horns on the head or thorax, making its body length greater than that of a female. Copulation is generally quick, but in some cases lasts for several hours. During copulation, sperm cells are transferred to the female to fertilize the egg.


title: Beetle

text: Essentially all beetles lay eggs, though some myrmecophilous Aleocharinae and some Chrysomelinae which live in mountains or the subarctic are ovoviviparous, laying eggs which hatch almost immediately. Beetle eggs generally have smooth surfaces and are soft, though the Cupedidae have hard eggs. Eggs vary widely between species: the eggs tend to be small in species with many instars (larval stages), and in those that lay large numbers of eggs.


title: Beetle

text: A female may lay from several dozen to several thousand eggs during her lifetime, depending on the extent of parental care. This ranges from the simple laying of eggs under a leaf, to the parental care provided by scarab beetles, which house, feed and protect their young. The Attelabidae roll leaves and lay their eggs inside the roll for protection.


title: Beetle

text: The larva is usually the principal feeding stage of the beetle life cycle. Larvae tend to feed voraciously once they emerge from their eggs. Some feed externally on plants, such as those of certain leaf beetles, while others feed within their food sources. Examples of internal feeders are most Buprestidae and longhorn beetles. The larvae of many beetle families are predatory like the adults (ground beetles, ladybirds, rove beetles). The larval period varies between species, but can be as long as several years. The larvae of skin beetles undergo a degree of reversed development when starved, and later grow back to the previously attained level of maturity. The cycle can be repeated many times (see Biological immortality). Larval morphology is highly varied amongst species, with well-developed and sclerotized heads, distinguishable thoracic and abdominal segments (usually the tenth, though sometimes the eighth or ninth).


title: Beetle

text: Beetle larvae can be differentiated from other insect larvae by their hardened, often darkened heads, the presence of chewing mouthparts, and spiracles along the sides of their bodies. Like adult beetles, the larvae are varied in appearance, particularly between beetle families. Beetles with somewhat flattened, highly mobile larvae include the ground beetles and rove beetles; their larvae are described as campodeiform. Some beetle larvae resemble hardened worms with dark head capsules and minute legs. These are elateriform larvae, and are found in the click beetle (Elateridae) and darkling beetle (Tenebrionidae) families. Some elateriform larvae of click beetles are known as wireworms. Beetles in the Scarabaeoidea have short, thick larvae described as scarabaeiform, more commonly known as grubs.


title: Beetle

text: All beetle larvae go through several instars, which are the developmental stages between each moult. In many species, the larvae simply increase in size with each successive instar as more food is consumed. In some cases, however, more dramatic changes occur. Among certain beetle families or genera, particularly those that exhibit parasitic lifestyles, the first instar (the planidium) is highly mobile to search out a host, while the following instars are more sedentary and remain on or within their host. This is known as hypermetamorphosis; it occurs in the Meloidae, Micromalthidae, and Ripiphoridae. The blister beetle "Epicauta vittata" (Meloidae), for example, has three distinct larval stages. Its first stage, the triungulin, has longer legs to go in search of the eggs of grasshoppers. After feeding for a week it moults to the second stage, called the caraboid stage, which resembles the larva of a carabid beetle. In another week it moults and assumes the appearance of a scarabaeid larva—the scarabaeidoid stage. Its penultimate larval stage is the pseudo-pupa or the coarcate larva, which will overwinter and pupate until the next spring.


title: Beetle

text: The larval period can vary widely. A fungus feeding staphylinid "Phanerota fasciata" undergoes three moults in 3.2 days at room temperature while "Anisotoma" sp. (Leiodidae) completes its larval stage in the fruiting body of slime mold in 2 days and possibly represents the fastest growing beetles. Dermestid beetles, "Trogoderma inclusum" can remain in an extended larval state under unfavourable conditions, even reducing their size between moults. A larva is reported to have survived for 3.5 years in an enclosed container.


title: Beetle

text: As with all endopterygotes, beetle larvae pupate, and from these pupae emerge fully formed, sexually mature adult beetles, or imagos. Pupae never have mandibles (they are adecticous). In most pupae, the appendages are not attached to the body and are said to be exarate; in a few beetles (Staphylinidae, Ptiliidae etc.) the appendages are fused with the body (termed as obtect pupae).


title: Beetle

text: Adults have extremely variable lifespans, from weeks to years, depending on the species. Some wood-boring beetles can have extremely long life-cycles. It is believed that when furniture or house timbers are infested by beetle larvae, the timber already contained the larvae when it was first sawn up. A birch bookcase 40 years old released adult "Eburia quadrigeminata" (Cerambycidae), while "Buprestis aurulenta" and other Buprestidae have been documented as emerging as much as 51 years after manufacture of wooden items.


title: Beetle

text: The elytra allow beetles to both fly and move through confined spaces, doing so by folding the delicate wings under the elytra while not flying, and folding their wings out just before takeoff. The unfolding and folding of the wings is operated by muscles attached to the wing base; as long as the tension on the radial and cubital veins remains, the wings remain straight. Some beetle species (many Cetoniinae; some Scarabaeinae, Curculionidae and Buprestidae) fly with the elytra closed, with the metathoracic wings extended under the lateral elytra margins. The altitude reached by beetles in flight varies. One study investigating the flight altitude of the ladybird species "Coccinella septempunctata" and "Harmonia axyridis" using radar showed that, whilst the majority in flight over a single location were at 150–195 m above ground level, some reached altitudes of over 1100 m.


title: Beetle

text: Many rove beetles have greatly reduced elytra, and while they are capable of flight, they most often move on the ground: their soft bodies and strong abdominal muscles make them flexible, easily able to wriggle into small cracks.


title: Beetle

text: Aquatic beetles use several techniques for retaining air beneath the water's surface. Diving beetles (Dytiscidae) hold air between the abdomen and the elytra when diving. Hydrophilidae have hairs on their under surface that retain a layer of air against their bodies. Adult crawling water beetles use both their elytra and their hind coxae (the basal segment of the back legs) in air retention, while whirligig beetles simply carry an air bubble down with them whenever they dive.


title: Beetle

text: Beetles have a variety of ways to communicate, including the use of pheromones. The mountain pine beetle emits a pheromone to attract other beetles to a tree. The mass of beetles are able to overcome the chemical defenses of the tree. After the tree's defenses have been exhausted, the beetles emit an anti-aggregation pheromone. This species can stridulate to communicate, but others may use sound to defend themselves when attacked.


title: Beetle

text: Parental care is found in a few families of beetle, perhaps for protection against adverse conditions and predators. The rove beetle "Bledius spectabilis" lives in salt marshes, so the eggs and larvae are endangered by the rising tide. The maternal beetle patrols the eggs and larvae, burrowing to keep them from flooding and asphyxiating, and protects them from the predatory carabid beetle "Dicheirotrichus gustavi" and from the parasitoidal wasp "Barycnemis blediator", which kills some 15% of the larvae.


title: Beetle

text: Burying beetles are attentive parents, and participate in cooperative care and feeding of their offspring. Both parents work to bury small animal carcass to serve as a food resource for their young and build a brood chamber around it. The parents prepare the carcass and protect it from competitors and from early decomposition. After their eggs hatch, the parents keep the larvae clean of fungus and bacteria and help the larvae feed by regurgitating food for them.


title: Beetle

text: Some dung beetles provide parental care, collecting herbivore dung and laying eggs within that food supply, an instance of mass provisioning. Some species do not leave after this stage, but remain to safeguard their offspring.


title: Beetle

text: Subsociality, where females guard their offspring, is well-documented in two families of Chrysomelidae, Cassidinae and Chrysomelinae.


title: Beetle

text: Eusociality involves cooperative brood care (including brood care of offspring from other individuals), overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. Few organisms outside Hymenoptera exhibit this behavior; the only beetle to do so is the weevil "Austroplatypus incompertus". This Australian species lives in horizontal networks of tunnels, in the heartwood of "Eucalyptus" trees. It is one of more than 300 species of wood-boring Ambrosia beetles which distribute the spores of ambrosia fungi. The fungi grow in the beetles' tunnels, providing food for the beetles and their larvae; female offspring remain in the tunnels and maintain the fungal growth, probably never reproducing. Cooperative brood care is also found in the bess beetles (Passalidae) where the larvae feed on the semi-digested faeces of the adults.


title: Beetle

text: Beetles are able to exploit a wide diversity of food sources available in their many habitats. Some are omnivores, eating both plants and animals. Other beetles are highly specialized in their diet. Many species of leaf beetles, longhorn beetles, and weevils are very host-specific, feeding on only a single species of plant. Ground beetles and rove beetles (Staphylinidae), among others, are primarily carnivorous and catch and consume many other arthropods and small prey, such as earthworms and snails. While most predatory beetles are generalists, a few species have more specific prey requirements or preferences. In some species, digestive ability relies upon a symbiotic relationship with fungi - some beetles have yeasts living their guts, including some yeasts previously undiscovered anywhere else.


title: Beetle

text: Decaying organic matter is a primary diet for many species. This can range from dung, which is consumed by coprophagous species (such as certain scarab beetles in the Scarabaeidae), to dead animals, which are eaten by necrophagous species (such as the carrion beetles, Silphidae). Some beetles found in dung and carrion are in fact predatory. These include members of the Histeridae and Silphidae, preying on the larvae of coprophagous and necrophagous insects. Many beetles feed under bark, some feed on wood while others feed on fungi growing on wood or leaf-litter. Some beetles have special mycangia, structures for the transport of fungal spores.


title: Beetle

text: Beetles, both adults and larvae, are the prey of many animal predators including mammals from bats to rodents, birds, lizards, amphibians, fishes, dragonflies, robberflies, reduviid bugs, ants, other beetles, and spiders. Beetles use a variety of anti-predator adaptations to defend themselves. These include camouflage and mimicry against predators that hunt by sight, toxicity, and defensive behaviour.


title: Beetle

text: Camouflage is common and widespread among beetle families, especially those that feed on wood or vegetation, such as leaf beetles (Chrysomelidae, which are often green) and weevils. In some species, sculpturing or various colored scales or hairs cause beetles such as the avocado weevil "Heilipus apiatus" to resemble bird dung or other inedible objects. Many beetles that live in sandy environments blend in with the coloration of that substrate.


title: Beetle

text: Some longhorn beetles (Cerambycidae) are effective Batesian mimics of wasps. Beetles may combine coloration with behavioural mimicry, acting like the wasps they already closely resemble. Many other beetles, including ladybirds, blister beetles, and lycid beetles secrete distasteful or toxic substances to make them unpalatable or poisonous, and are often aposematic, where bright or contrasting coloration warn off predators; many beetles and other insects mimic these chemically protected species.


title: Beetle

text: Chemical defense is important in some species, usually being advertised by bright aposematic colors. Some Tenebrionidae use their posture for releasing noxious chemicals to warn off predators. Chemical defenses may serve purposes other than just protection from vertebrates, such as protection from a wide range of microbes. Some species sequester chemicals from the plants they feed on, incorporating them into their own defenses.


title: Beetle

text: Other species have special glands to produce deterrent chemicals. The defensive glands of carabid ground beetles produce a variety of hydrocarbons, aldehydes, phenols, quinones, esters, and acids released from an opening at the end of the abdomen. African carabid beetles (for example, "Anthia") employ the same chemicals as ants: formic acid. Bombardier beetles have well-developed pygidial glands that empty from the sides of the intersegment membranes between the seventh and eighth abdominal segments. The gland is made of two containing chambers, one for hydroquinones and hydrogen peroxide, the other holding hydrogen peroxide and catalase enzymes. These chemicals mix and result in an explosive ejection, reaching a temperature of around , with the breakdown of hydroquinone to hydrogen, oxygen, and quinone. The oxygen propels the noxious chemical spray as a jet that can be aimed accurately at predators.


title: Beetle

text: Large ground-dwelling beetles such as Carabidae, the rhinoceros beetle and the longhorn beetles defend themselves using strong mandibles, or heavily sclerotised (armored) spines or horns to deter or fight off predators. Many species of weevil that feed out in the open on leaves of plants react to attack by employing a drop-off reflex. Some combine it with thanatosis, in which they close up their appendages and "play dead". The click beetles (Elateridae) can suddenly catapult themselves out of danger by releasing the energy stored by a click mechanism, which consists of a stout spine on the prosternum and a matching groove in the mesosternum. Some species startle an attacker by producing sounds through a process known as stridulation.


title: Beetle

text: A few species of beetles are ectoparasitic on mammals. One such species, "Platypsyllus castoris", parasitises beavers ("Castor" spp.). This beetle lives as a parasite both as a larva and as an adult, feeding on epidermal tissue and possibly on skin secretions and wound exudates. They are strikingly flattened dorsoventrally, no doubt as an adaptation for slipping between the beavers' hairs. They are wingless and eyeless, as are many other ectoparasites. Others are kleptoparasites of other invertebrates, such as the small hive beetle ("Aethina tumida") that infests honey bee nests, while many species are parasitic inquilines or commensal in the nests of ants. A few groups of beetles are primary parasitoids of other insects, feeding off of, and eventually killing their hosts.


title: Beetle

text: Beetle-pollinated flowers are usually large, greenish or off-white in color, and heavily scented. Scents may be spicy, fruity, or similar to decaying organic material. Beetles were most likely the first insects to pollinate flowers. Most beetle-pollinated flowers are flattened or dish-shaped, with pollen easily accessible, although they may include traps to keep the beetle longer. The plants' ovaries are usually well protected from the biting mouthparts of their pollinators. The beetle families that habitually pollinate flowers are the Buprestidae, Cantharidae, Cerambycidae, Cleridae, Dermestidae, Lycidae, Melyridae, Mordellidae, Nitidulidae and Scarabaeidae. Beetles may be particularly important in some parts of the world such as semiarid areas of southern Africa and southern California and the montane grasslands of KwaZulu-Natal in South Africa.


title: Beetle

text: Mutualism is well known in a few beetles, such as the ambrosia beetle, which partners with fungi to digest the wood of dead trees. The beetles excavate tunnels in dead trees in which they cultivate fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases spores of its fungal symbiont. The fungus penetrates the plant's xylem tissue, digests it, and concentrates the nutrients on and near the surface of the beetle gallery, so the weevils and the fungus both benefit. The beetles cannot eat the wood due to toxins, and uses its relationship with fungi to help overcome the defenses of its host tree in order to provide nutrition for their larvae. Chemically mediated by a bacterially produced polyunsaturated peroxide, this mutualistic relationship between the beetle and the fungus is coevolved.


title: Beetle

text: About 90% of beetle species enter a period of adult diapause, a quiet phase with reduced metabolism to tide unfavourable environmental conditions. Adult diapause is the most common form of diapause in Coleoptera. To endure the period without food (often lasting many months) adults prepare by accumulating reserves of lipids, glycogen, proteins and other substances needed for resistance to future hazardous changes of environmental conditions. This diapause is induced by signals heralding the arrival of the unfavourable season; usually the cue is photoperiodic. Short (decreasing) day length serves as a signal of approaching winter and induces winter diapause (hibernation). A study of hibernation in the Arctic beetle "Pterostichus brevicornis" showed that the body fat levels of adults were highest in autumn with the alimentary canal filled with food, but empty by the end of January. This loss of body fat was a gradual process, occurring in combination with dehydration.


title: Beetle

text: All insects are poikilothermic, so the ability of a few beetles to live in extreme environments depends on their resilience to unusually high or low temperatures. The bark beetle "Pityogenes chalcographus" can survive whilst overwintering beneath tree bark; the Alaskan beetle "Cucujus clavipes puniceus" is able to withstand ; its larvae may survive . At these low temperatures, the formation of ice crystals in internal fluids is the biggest threat to survival to beetles, but this is prevented through the production of antifreeze proteins that stop water molecules from grouping together. The low temperatures experienced by "Cucujus clavipes" can be survived through their deliberate dehydration in conjunction with the antifreeze proteins. This concentrates the antifreezes several fold. The hemolymph of the mealworm beetle "Tenebrio molitor" contains several antifreeze proteins. The Alaskan beetle "Upis ceramboides" can survive −60 °C: its cryoprotectants are xylomannan, a molecule consisting of a sugar bound to a fatty acid, and the sugar-alcohol, threitol.


title: Beetle

text: Conversely, desert dwelling beetles are adapted to tolerate high temperatures. For example, the Tenebrionid beetle "Onymacris rugatipennis" can withstand . Tiger beetles in hot, sandy areas are often whitish (for example, "Habroscelimorpha dorsalis"), to reflect more heat than a darker color would. These beetles also exhibits behavioural adaptions to tolerate the heat: they are able to stand erect on their tarsi to hold their bodies away from the hot ground, seek shade, and turn to face the sun so that only the front parts of their heads are directly exposed.


title: Beetle

text: The fogstand beetle of the Namib Desert, "Stenocara gracilipes", is able to collect water from fog, as its elytra have a textured surface combining hydrophilic (water-loving) bumps and waxy, hydrophobic troughs. The beetle faces the early morning breeze, holding up its abdomen; droplets condense on the elytra and run along ridges towards their mouthparts. Similar adaptations are found in several other Namib desert beetles such as "Onymacris unguicularis".


title: Beetle

text: Some terrestrial beetles that exploit shoreline and floodplain habitats have physiological adaptations for surviving floods. In the event of flooding, adult beetles may be mobile enough to move away from flooding, but larvae and pupa often cannot. Adults of "Cicindela togata" are unable to survive immersion in water, but larvae are able to survive a prolonged period, up to 6 days, of anoxia during floods. Anoxia tolerance in the larvae may have been sustained by switching to anaerobic metabolic pathways or by reducing metabolic rate. Anoxia tolerance in the adult carabid beetle "Pelophilia borealis" was tested in laboratory conditions and it was found that they could survive a continuous period of up to 127 days in an atmosphere of 99.9% nitrogen at 0 °C.


title: Beetle

text: Many beetle species undertake annual mass movements which are termed as migrations. These include the pollen beetle "Meligethes aeneus" and many species of coccinellids. These mass movements may also be opportunistic, in search of food, rather than seasonal. A 2008 study of an unusually large outbreak of Mountain Pine Beetle ("Dendroctonus ponderosae") in British Columbia found that beetles were capable of flying 30–110 km per day in densities of up to 18,600 beetles per hectare.


title: Beetle

text: Several species of dung beetle, especially the sacred scarab, "Scarabaeus sacer", were revered in Ancient Egypt. The hieroglyphic image of the beetle may have had existential, fictional, or ontologic significance. Images of the scarab in bone, ivory, stone, Egyptian faience, and precious metals are known from the Sixth Dynasty and up to the period of Roman rule. The scarab was of prime significance in the funerary cult of ancient Egypt. The scarab was linked to Khepri, the god of the rising sun, from the supposed resemblance of the rolling of the dung ball by the beetle to the rolling of the sun by the god. Some of ancient Egypt's neighbors adopted the scarab motif for seals of varying types. The best-known of these are the Judean LMLK seals, where eight of 21 designs contained scarab beetles, which were used exclusively to stamp impressions on storage jars during the reign of Hezekiah. Beetles are mentioned as a symbol of the sun, as in ancient Egypt, in Plutarch's 1st century "Moralia". The Greek Magical Papyri of the 2nd century BC to the 5th century AD describe scarabs as an ingredient in a spell.


title: Beetle

text: Pliny the Elder discusses beetles in his "Natural History", describing the stag beetle: "Some insects, for the preservation of their wings, are covered with (elytra)—the beetle, for instance, the wing of which is peculiarly fine and frail. To these insects a sting has been denied by Nature; but in one large kind we find horns of a remarkable length, two-pronged at the extremities, and forming pincers, which the animal closes when it is its intention to bite." The stag beetle is recorded in a Greek myth by Nicander and recalled by Antoninus Liberalis in which Cerambus is turned into a beetle: "He can be seen on trunks and has hook-teeth, ever moving his jaws together. He is black, long and has hard wings like a great dung beetle". The story concludes with the comment that the beetles were used as toys by young boys, and that the head was removed and worn as a pendant.


title: Beetle

text: About 75% of beetle species are phytophagous in both the larval and adult stages. Many feed on economically important plants and stored plant products, including trees, cereals, tobacco, and dried fruits. Some, such as the boll weevil, which feeds on cotton buds and flowers, can cause extremely serious damage to agriculture. The boll weevil crossed the Rio Grande near Brownsville, Texas, to enter the United States from Mexico around 1892, and had reached southeastern Alabama by 1915. By the mid-1920s, it had entered all cotton-growing regions in the US, traveling per year. It remains the most destructive cotton pest in North America. Mississippi State University has estimated, since the boll weevil entered the United States, it has cost cotton producers about $13 billion, and in recent times about $300 million per year.


title: Beetle

text: The bark beetle, elm leaf beetle and the Asian longhorned beetle ("Anoplophora glabripennis") are among the species that attack elm trees. Bark beetles (Scolytidae) carry Dutch elm disease as they move from infected breeding sites to healthy trees. The disease has devastated elm trees across Europe and North America.


title: Beetle

text: Some species of beetle have evolved immunity to insecticides. For example, the Colorado potato beetle, "Leptinotarsa decemlineata", is a destructive pest of potato plants. Its hosts include other members of the Solanaceae, such as nightshade, tomato, eggplant and capsicum, as well as the potato. Different populations have between them developed resistance to all major classes of insecticide. The Colorado potato beetle was evaluated as a tool of entomological warfare during World War II, the idea being to use the beetle and its larvae to damage the crops of enemy nations. Germany tested its Colorado potato beetle weaponisation program south of Frankfurt, releasing 54,000 beetles.


title: Beetle

text: The death watch beetle, "Xestobium rufovillosum" (Ptinidae), is a serious pest of older wooden buildings in Europe. It attacks hardwoods such as oak and chestnut, always where some fungal decay has taken or is taking place. The actual introduction of the pest into buildings is thought to take place at the time of construction.


title: Beetle

text: Other pests include the coconut hispine beetle, "Brontispa longissima", which feeds on young leaves, seedlings and mature coconut trees, causing serious economic damage in the Philippines. The mountain pine beetle is a destructive pest of mature or weakened lodgepole pine, sometimes affecting large areas of Canada.


title: Beetle

text: Beetles can be beneficial to human economics by controlling the populations of pests. The larvae and adults of some species of lady beetles (Coccinellidae) feed on aphids that are pests. Other lady beetles feed on scale insects, whitefly and mealybugs. If normal food sources are scarce, they may feed on small caterpillars, young plant bugs, or honeydew and nectar. Ground beetles (Carabidae) are common predators of many insect pests, including fly eggs, caterpillars, and wireworms. Ground beetles can help to control weeds by eating their seeds in the soil, reducing the need for herbicides to protect crops. The effectiveness of some species in reducing certain plant populations has resulted in the deliberate introduction of beetles in order to control weeds. For example, the genus "Zygogramma" is native to North America but has been used to control "Parthenium hysterophorus" in India and "Ambrosia artemisiifolia" in Russia.


title: Beetle

text: Dung beetles (Scarabidae) have been successfully used to reduce the populations of pestilent flies, such as "Musca vetustissima" and "Haematobia exigua" which are serious pests of cattle in Australia. The beetles make the dung unavailable to breeding pests by quickly rolling and burying it in the soil, with the added effect of improving soil fertility, tilth, and nutrient cycling. The Australian Dung Beetle Project (1965–1985), introduced species of dung beetle to Australia from South Africa and Europe to reduce populations of "Musca vetustissima", following successful trials of this technique in Hawaii. The American Institute of Biological Sciences reports that dung beetles save the United States cattle industry an estimated US$380 million annually through burying above-ground livestock feces.


title: Beetle

text: The Dermestidae are often used in taxidermy and in the preparation of scientific specimens, to clean soft tissue from bones. Larvae feed on and remove cartilage along with other soft tissue.


title: Beetle

text: Beetles are the most widely eaten insects, with about 344 species used as food, usually at the larval stage. The mealworm (the larva of the darkling beetle) and the rhinoceros beetle are among the species commonly eaten. A wide range of species is also used in folk medicine to treat those suffering from a variety of disorders and illnesses, though this is done without clinical studies supporting the efficacy of such treatments.


title: Beetle

text: Due to their habitat specificity, many species of beetles have been suggested as suitable as indicators, their presence, numbers, or absence providing a measure of habitat quality. Predatory beetles such as the tiger beetles (Cicindelidae) have found scientific use as an indicator taxon for measuring regional patterns of biodiversity. They are suitable for this as their taxonomy is stable; their life history is well described; they are large and simple to observe when visiting a site; they occur around the world in many habitats, with species specialised to particular habitats; and their occurrence by species accurately indicates other species, both vertebrate and invertebrate. According to the habitats, many other groups such as the rove beetles in human-modified habitats, dung beetles in savannas and saproxylic beetles in forests have been suggested as potential indicator species.


title: Beetle

text: Many beetles have durable elytra that has been used as material in art, with beetlewing the best example. Sometimes, they are incorporated into ritual objects for their religious significance. Whole beetles, either as-is or encased in clear plastic, are made into objects ranging from cheap souvenirs such as key chains to expensive fine-art jewellery. In parts of Mexico, beetles of the genus "Zopherus" are made into living brooches by attaching costume jewelry and golden chains, which is made possible by the incredibly hard elytra and sedentary habits of the genus.


title: Beetle

text: Fighting beetles are used for entertainment and gambling. This sport exploits the territorial behavior and mating competition of certain species of large beetles. In the Chiang Mai district of northern Thailand, male "Xylotrupes" rhinoceros beetles are caught in the wild and trained for fighting. Females are held inside a log to stimulate the fighting males with their pheromones. These fights may be competitive and involve gambling both money and property. In South Korea the Dytiscidae species "Cybister tripunctatus" is used in a roulette-like game.


title: Beetle

text: Beetles are sometimes used as instruments: the Onabasulu of Papua New Guinea historically used the ""hugu"" weevil "Rhynchophorus ferrugineus" as a musical instrument by letting the human mouth serve as a variable resonance chamber for the wing vibrations of the live adult beetle.


title: Beetle

text: Some species of beetle are kept as pets, for example diving beetles (Dytiscidae) may be kept in a domestic fresh water tank.


title: Beetle

text: In Japan the practice of keeping horned rhinoceros beetles (Dynastinae) and stag beetles (Lucanidae) is particularly popular amongst young boys. Such is the popularity in Japan that vending machines dispensing live beetles were developed in 1999, each holding up to 100 stag beetles.


title: Beetle

text: Beetle collecting became extremely popular in the Victorian era. The naturalist Alfred Russel Wallace collected (by his own count) a total of 83,200 beetles during the eight years described in his 1869 book "The Malay Archipelago", including 2,000 species new to science.


title: Beetle

text: Several coleopteran adaptations have attracted interest in biomimetics with possible commercial applications. The bombardier beetle's powerful repellent spray has inspired the development of a fine mist spray technology, claimed to have a low carbon impact compared to aerosol sprays. Moisture harvesting behavior by the Namib desert beetle ("Stenocara gracilipes") has inspired a self-filling water bottle which utilises hydrophilic and hydrophobic materials to benefit people living in dry regions with no regular rainfall.


title: Beetle

text: Living beetles have been used as cyborgs. A Defense Advanced Research Projects Agency funded project implanted electrodes into "Mecynorhina torquata" beetles, allowing them to be remotely controlled via a radio receiver held on its back, as proof-of-concept for surveillance work. Similar technology has been applied to enable a human operator to control the free-flight steering and walking gaits of "Mecynorhina torquata" as well as graded turning and backward walking of "Zophobas morio".


title: Beetle

text: Research published in 2020 sought to create a robotic camera backpack for beetles. Miniature cameras weighing 248 mg were attached to live beetles of the Tenebrionid genera "Asbolus" and "Eleodes". The cameras filmed over a 60° range for up to 6 hours.


title: Beetle

text: Since beetles form such a large part of the world's biodiversity, their conservation is important, and equally, loss of habitat and biodiversity is essentially certain to impact on beetles. Many species of beetles have very specific habitats and long life cycles that make them vulnerable. Some species are highly threatened while others are already feared extinct. Island species tend to be more susceptible as in the case of "Helictopleurus undatus" of Madagascar which is thought to have gone extinct during the late 20th century. Conservationists have attempted to arouse a liking for beetles with flagship species like the stag beetle, "Lucanus cervus", and tiger beetles (Cicindelidae). In Japan the Genji firefly, "Luciola cruciata", is extremely popular, and in South Africa the Addo elephant dung beetle offers promise for broadening ecotourism beyond the big five tourist mammal species. Popular dislike of pest beetles, too, can be turned into public interest in insects, as can unusual ecological adaptations of species like the fairy shrimp hunting beetle, "Cicinis bruchi".


title: Operation Entebbe

text: Operation Entebbe, also known as the Entebbe Raid or Operation Thunderbolt, was a counter-terrorist hostage-rescue mission carried out by commandos of the Israel Defense Forces (IDF) at Entebbe Airport in Uganda on 4 July 1976.


title: Operation Entebbe

text: A week earlier, on 27 June, an Air France Airbus A300 jet airliner with 248 passengers had been hijacked by two members of the Popular Front for the Liberation of Palestine – External Operations (PFLP-EO) under orders of Wadie Haddad (who had earlier broken away from the PFLP of George Habash), and two members of the German Revolutionary Cells. The hijackers had the stated objective to free 40 Palestinian and affiliated militants imprisoned in Israel and 13 prisoners in four other countries in exchange for the hostages. The flight, which had originated in Tel Aviv with the destination of Paris, was diverted after a stopover in Athens via Benghazi to Entebbe, the main airport of Uganda. The Ugandan government supported the hijackers, and dictator Idi Amin, who had been informed of the hijacking from the beginning, personally welcomed them. After moving all hostages from the aircraft to a disused airport building, the hijackers separated all Israelis and several non-Israeli Jews from the larger group and forced them into a separate room. Over the following two days, 148 non-Israeli hostages were released and flown out to Paris. Ninety-four, mainly Israeli, passengers along with the 12-member Air France crew, remained as hostages and were threatened with death.


title: Operation Entebbe

text: The IDF acted on information provided by the Israeli intelligence agency Mossad. The hijackers threatened to kill the hostages if their prisoner release demands were not met. This threat led to the planning of the rescue operation. These plans included preparation for armed resistance from the Uganda Army.


title: Operation Entebbe

text: The operation took place at night. Israeli transport planes carried 100 commandos over to Uganda for the rescue operation. The operation, which took a week of planning, lasted 90 minutes. Of the 106 remaining hostages, 102 were rescued and three were killed. The other hostage was in a hospital and was later killed. Five Israeli commandos were wounded, and one, unit commander Lt. Col. Yonatan Netanyahu, was killed. Netanyahu was the older brother of Benjamin Netanyahu, who would later become Prime Minister of Israel. All the hijackers and forty-five Ugandan soldiers were killed, and eleven Soviet-built MiG-17s and MiG-21s of Uganda's air force were destroyed. Kenyan sources supported Israel, and in the aftermath of the operation, Idi Amin issued orders to retaliate and kill Kenyans present in Uganda. As a result, 245 Kenyans in Uganda were killed and 3,000 fled the country.


title: Operation Entebbe

text: Operation Entebbe, which had the military codename Operation Thunderbolt, is sometimes referred to retroactively as Operation Jonathan in memory of the unit's leader, Yonatan Netanyahu.


title: Operation Entebbe

text: On 27 June 1976, Air France Flight 139, an Airbus A300B4-203, registration (c/n 019), departed from Tel Aviv, Israel, carrying 246 mainly Jewish and Israeli passengers and a crew of 12. The plane flew to Athens, Greece, where it picked up an additional 58 passengers, including four hijackers. It departed for Paris at 12:30 pm. Just after takeoff, the flight was hijacked by two Palestinians from the Popular Front for the Liberation of Palestine – External Operations (PFLP-EO), and by two Germans, Wilfried Böse and Brigitte Kuhlmann, from the German Revolutionary Cells. The hijackers diverted the flight to Benghazi, Libya. There it was held on the ground for seven hours for refuelling. During that time the hijackers released British-born Israeli citizen Patricia Martell, who pretended to have a miscarriage. The plane left Benghazi and at 3:15 pm on the 28th, more than 24 hours after the flight's original departure, it arrived at Entebbe Airport in Uganda.


title: Operation Entebbe

text: At Entebbe, the four hijackers were joined by at least four others, supported by the forces of Uganda's president, Idi Amin. The hijackers transferred the passengers to the transit hall of the disused former airport terminal where they kept them under guard for the following days. Amin came to visit the hostages almost on a daily basis, updating them on developments and promising to use his efforts to have them freed through negotiations.


title: Operation Entebbe

text: On 28 June, a PFLP-EO hijacker issued a declaration and formulated their demands: In addition to a ransom of US$5 million for the release of the airplane, they demanded the release of 53 Palestinian and pro-Palestinian militants, 40 of whom were prisoners in Israel. They threatened that if these demands were not met, they would begin to kill hostages on 1 July 1976.


title: Operation Entebbe

text: On 29 June, after Ugandan soldiers had opened an entrance to a room next to the crowded waiting hall by destroying a separating wall, the hijackers separated the Israelis (including those holding dual citizenship) from the other hostages and told them to move to the adjoining room. As they did so, a Holocaust survivor showed hijacker Wilfried Böse a camp registration number tattooed on his arm. Böse protested "I'm no Nazi! ... I am an idealist". In addition, five non-Israeli hostages – two ultra-orthodox Jewish couples from the US and Belgium and a French resident of Israel – were forced to join the Israeli group. According to Monique Epstein Khalepski, the French hostage among the five, the captors had singled them out for questioning and suspected them of hiding their Israeli identities. On the other hand, according to French hostage Michel Cojot-Goldberg, the captors failed to identify at least one Israeli among the passengers who was a military officer with dual citizenship then using his non-Israeli passport and he was later freed as part of the second release of non-Israeli hostages. US citizen Janet Almog, Frenchwoman Jocelyne Monier (whose husband or boyfriend was Israeli), and French-Israeli dual citizen Jean-Jacques Mimouni, whose name had not been called up during the reading of the original passport-based list, reportedly joined the Israeli hostage group by their own choice.


title: Operation Entebbe

text: On 30 June, the hijackers released 48 hostages. The released were picked from among the non-Israeli group – mainly elderly and sick passengers and mothers with children. Forty-seven of them were flown by a chartered Air France Boeing 747 out of Entebbe to Paris, and one passenger was treated in hospital for a day. On 1 July, after the Israeli government had conveyed its agreement to negotiations, the hostage-takers extended their deadline to noon on 4 July and released another group of 100 non-Israeli captives who again were flown to Paris a few hours later. Among the 106 hostages staying behind with their captors at Entebbe airport were the 12 members of the Air France crew who refused to leave, about ten young French passengers, and the Israeli group of some 84 people.


title: Operation Entebbe

text: In the week before the raid, Israel tried using political avenues to obtain the release of the hostages. Many sources indicate that the Israeli cabinet was prepared to release Palestinian prisoners if a military solution seemed unlikely to succeed. A retired IDF officer, Baruch "Burka" Bar-Lev, had known Idi Amin for many years and was considered to have a strong personal relationship with him. At the request of the cabinet, he spoke with Amin on the phone many times, trying to gain the release of the hostages, without success. The Israeli government also approached the United States government to deliver a message to Egyptian president Anwar Sadat, asking him to request that Amin release the hostages. Prime minister Yitzhak Rabin and defence minister Shimon Peres spent one week disagreeing on whether to give in to the hijackers' demands (Rabin's position) or not, to prevent more terrorism (Peres' position).


title: Operation Entebbe

text: At the 1 July deadline, the Israeli cabinet offered to negotiate with the hijackers to extend the deadline to 4 July. Amin also asked them to extend the deadline until that date. This meant he could take a diplomatic trip to Port Louis, Mauritius, to officially hand over chairmanship of the Organisation of African Unity to Seewoosagur Ramgoolam. This extension of the hostage deadline proved crucial to providing Israeli forces enough time to get to Entebbe.


title: Operation Entebbe

text: On 3 July, at 18:30, the Israeli cabinet approved a rescue mission, presented by Major General Yekutiel Adam and Brigadier General Dan Shomron. Shomron was appointed as the operation commander.