title: Ron Francis

text: Ronald Michael Francis Jr. (born March 1, 1963) is a Canadian former professional ice hockey centre. He is the general manager of the Seattle Kraken that inaugurated its first season in 2021. Drafted fourth overall in the 1981 NHL Entry Draft, Francis played 23 seasons in the NHL for the Hartford Whalers, Pittsburgh Penguins, Carolina Hurricanes, and Toronto Maple Leafs. Upon retiring from professional ice hockey in 2004, Francis stood second all-time in career assists (1,249), behind only Wayne Gretzky; fifth in career points (1,798); third in games played (1,731); and 27th in career goals (549).


title: Ron Francis

text: In 2014, Francis was named as the general manager for the Hurricanes, replacing Jim Rutherford, who had been with the franchise ever since the team's move to Raleigh, North Carolina. Two years before, Francis had become a minority owner of the team as part of the five-man investor group, Playmakers Management. In March 2018, he was promoted to the president of hockey operations role. His Hurricanes contract was terminated on April 30, 2018. On July 18, 2019, he was hired as the first general manager of the Seattle Kraken.


title: Ron Francis

text: As a youth, Francis played in the 1975 and 1976 Quebec International Pee-Wee Hockey Tournaments with a minor ice hockey team from Sault Ste. Marie, Ontario.


title: Ron Francis

text: Francis was drafted by the Hartford Whalers in the first round, fourth overall, of the 1981 NHL Entry Draft. He was a model of consistency and durability, averaging more than a point a game in over 1,700 games in 23 seasons, and (not counting the lockout-shortened 1994–95 season) averaging just under 77 games played a season. His three Lady Byng Trophies attest to his gentlemanly conduct on and off the ice. Francis stands second all-time in career assists behind Wayne Gretzky with 1,249, fifth in career points (1,798), third in games played (1,731), and twenty-sixth in career goals (549).


title: Ron Francis

text: Francis played almost ten seasons with the Whalers, serving as captain for almost six and setting nearly every offensive record in franchise history. He was traded to the Pittsburgh Penguins on March 4, 1991 with Ulf Samuelsson and Grant Jennings, in exchange for Jeff Parker, Zarley Zalapski, and John Cullen. The trade became a coup for Pittsburgh, where he centred a formidable second line behind Mario Lemieux, as the Penguins won their first Stanley Cup less than three months later. The threat posed by the Francis-centered second line, formidable in its own right, often required opposing teams to throw their best defenseman against two lines, instead of zeroing in on Lemieux's line, thus creating more opportunities for both top lines. Francis was also known as one of the best face-off men in the NHL in this period, in both the offensive and defensive ends. In power-play and empty-net situations, his key face-off wins often led to Penguin goals.


title: Ron Francis

text: Francis was indispensable the following year, as Pittsburgh repeated as champions, in leading the team during the absence of Lemieux in the 1992 playoffs – and in scoring the Cup-clinching goal against the Chicago Blackhawks. At the same time, the trade that brought him to Pittsburgh is considered to be one of the most one-sided trades in NHL history (though "The Hockey News" suggested that Hartford had gotten the better end of the trade at the time); the players Hartford acquired never approached the numbers or impact Francis produced there or with Pittsburgh. Francis would spend seven seasons in Pittsburgh, captaining the team twice, and becoming the first Penguin to win the Selke Trophy in 1995.


title: Ron Francis

text: Francis returned to his original organization as a free agent for 1998–99, signing with the Carolina Hurricanes (who had moved from Hartford the previous season). He spent the next 5.5 seasons padding his franchise records. He still ranks first all-time in Whalers/Hurricanes history in points, goals, assists and games played. At the time of his retirement, his 1,175 points were more than double those of then-runner up Kevin Dineen. He captained the Hurricanes to a surprise appearance in the 2002 Stanley Cup Finals and scored the winning goal for the Hurricanes in overtime of Game 1, before losing to the Detroit Red Wings in five games.


title: Ron Francis

text: Francis finished his career with a brief stint with the Toronto Maple Leafs, traded there by the Hurricanes in March 2004 to allow him one last run at the Stanley Cup. He retired from the NHL before the 2005–06 season and assumed a position with the Raleigh Youth Hockey Association.


title: Ron Francis

text: In June 2011, Francis assumed the position of director of hockey operations with the Carolina Hurricanes before later being named general manager of the team in 2014. On March 7, 2018, Francis was named president of hockey operations by new Hurricanes' majority owner Thomas Dundon. His Hurricanes contract was terminated on April 30, 2018. On July 18, 2019, Francis was named the first general manager of the Seattle Kraken.


title: Ron Francis

text: Francis is married to the former Mary Lou Robie, a native of Stamford, Connecticut whom he met in Hartford during his tenure with the Whalers. They married in 1986 and have three children: Kaitlyn (b. 1991), Michael (b. 1993), and Connor (b. 1996). Francis is considered a popular sports figure in Hartford, Pittsburgh and Raleigh, and is also noted for his humanitarian and charity work. Francis also has the distinction of being the first ice hockey player inducted into the North Carolina Sports Hall of Fame.


title: Ron Francis

text: Francis won two Stanley Cups, in 1990–91 and the following season, with the Pittsburgh Penguins. Statistically, his best season was 1995–96, when he recorded 119 points; that season, he led the NHL in assists, with 92. The previous season, he had not only led the League in assists with 48 over the lockout-shortened, half-season schedule, but became the first player to win both the Frank J. Selke Trophy and the Lady Byng Trophy in the same season.


title: Ron Francis

text: Francis' Whalers number 10 jersey was raised at the Hartford Civic Center on January 6, 2006 (though not officially retired, the Whalers organization no longer existing in Hartford to retire it), along with Ulf Samuelsson's number 5 and Kevin Dineen's number 11. Additionally, his Hurricanes number 10 jersey was retired by the Carolina organization on January 28, 2006. He was also pictured in the Pittsburgh Penguins Ring of Honor that formerly circled the upper level of the Pittsburgh Civic Arena.


title: Ron Francis

text: On June 28, 2007, Francis was selected to enter the Hockey Hall of Fame in his first year on the ballot. He was formally inducted on November 12, 2007.


title: OSV-96

text: OSV-96 (Russian: ОСВ-96) is a Russian heavy semi-automatic precision rifle chambered for the 12.7×108mm.


title: OSV-96

text: The rifle is capable of engaging infantry at a distance of up to 1800 meters and can combat material targets at ranges up to 2500 meters. As an effective anti-sniper weapon, OSV-96 keeps the shooter outside of the effective range of conventional calibers providing a distinct advantage over lower caliber rifles. Specialized high-accuracy 12.7×108mm 7N34 59.0 gram FMJ and AP sniper cartridges have been developed for Russian .50-caliber sniper rifles such as this rifle. Large caliber machine gun cartridges can also be used for firing, but with limited accuracy. With 7N34 ammunition, its accuracy is claimed as 1.5 MOA at 100 meters or better, equal to the Barrett M107.


title: OSV-96

text: The OSV-96 folds in half in between the barrel/chamber and receiver compartments to shorten its length for ease of transportation. The rifle features a free-floating barrel in combination with a large muzzle brake to greatly decrease its recoil.


title: Krishna Hutheesing

text: Krishna Nehru Hutheesing (2 November 1907 – 9 November 1967) was an Indian writer, the youngest sister of Jawaharlal Nehru and Vijaya Lakshmi Pandit, and part of the Nehru-Gandhi family.


title: Krishna Hutheesing

text: Born Krishna Nehru, in Mirganj, Allahabad to Motilal Nehru, an Indian independence activist and leader of the Indian National Congress, and Swarup Rani, she was married to Gunottam (Raja) Hutheesing, who belonged to an Ahmedabad Jain family that built the Hutheesing Jain Temple. During the later 1950s, he became critic of Nehru and in 1959, supported former Governor General C. Rajagopalachari, to form a conservative market liberal political party known as the Swatantra Party.


title: Krishna Hutheesing

text: She and her husband fought for India's independence and spent a great deal of time in jail. Raja's terms in jail came while they were raising their two young sons, Harsha Hutheesing and Ajit Hutheesing.


title: Krishna Hutheesing

text: In 1950, Krishna and her husband toured the United States on a lecture tour. In late May 1958 Krishna spent three days in Israel. Her host was Yigal Alon, who a year earlier founded 'The Israel-India Friendship League' as a tool to circumvent the then Indian government policy to avoid direct diplomatic relations between the two states.


title: Krishna Hutheesing

text: Mrs. Hutheesing documented her life as well as the lives of her brother, Jawaharlal and her niece, Indira Gandhi, in a series of books that intertwine history with personal anecdotes including "We Nehrus," "With No Regrets- An Autobiography," and "Dear to Behold: An Intimate Portrait of Indira Gandhi"


title: Krishna Hutheesing

text: Her husband, Raja Hutheesing, also wrote books: "The Great Peace: An Asian's Candid Report on Red China" (1953), "Window on China" (1953), and "Tibet fights for freedom : the story of the March 1959 uprising" (1960).


title: Krishna Hutheesing

text: Mrs. Hutheesing was associated with the 'Voice of America' and gave several talks. She died in London in 1967.


title: Runa Laila

text: Runa Laila (, ; born 17 November 1952) is a Bangladeshi playback singer and composer. She started her career in Pakistan film industry in the late 1960s. Her style of singing is inspired by Pakistani playback singer Ahmed Rushdi and she also made a pair with him after replacing another singer Mala. Her playback singing in films – "The Rain" (1976), "Jadur Banshi" (1977), "Accident" (1989), "Ontore Ontore" (1994), "Devdas" (2013) and "Priya Tumi Shukhi Hou" (2014) - earned her seven Bangladesh National Film Awards for Best Female Playback Singer. She won the Best Music Composer award for the film "Ekti Cinemar Golpo" (2018).


title: Runa Laila

text: Laila was born in Sylhet to Syed Mohammed Imdad Ali, a civil servant posted in Karachi, and Amina Laila. She started taking dance lessons of Kathak and Bharatanatyam genre. In those days, Ahmed Rushdi was the leading film singer who introduced rock n roll, disco and other modern genres to South Asian music. Following Rushdi's success, Christian bands specialising in jazz started performing at various night clubs and hotel lobbies in Karachi, Hyderabad, Mumbai, Dhaka and Lahore. Laila became a fan of singer Ahmed Rushdi whom she considered her guru (teacher), and tried to emulate not only his singing style but also the way he used to perform on the stage. She then learned classical music with her elder sister Dina Laila (d. 1976).


title: Runa Laila

text: While she was a student of Saint Lawrence Convent, she won an inter-school singing competition in Karachi in the then West Pakistan. She, along with her sister, were trained by Ustad Abdul Kader Peyarang and Ustad Habibuddin Ahmed. Her cousin, Anjumara Begum, had already been a known singer. When Laila was 12, she performed as a playback singer for a male child actor in the Urdu language film "Jugnu". The song was titled "Gudia Si Munni Meri".


title: Runa Laila

text: In 1966, Laila made her breakthrough in the Pakistani film industry with the song "Unki Nazron Sey Mohabbat Ka Jo Paigham Mila" for the Urdu film "Hum Dono". She used to perform on PTV. In PTV, she had a show called "Bazm E Laila". She started appearing on the "Zia Mohyuddin Show" (1972–74) and later sang songs for films in the 1970s such as the film Umrao Jaan Ada (1972).


title: Runa Laila

text: Laila moved to Bangladesh along with her family in 1974. Her first Bengali song was "O Amar Jibon Shathi" for the film Jibon Shathi (1976), composed by Satya Saha. Shortly after had her first concert in India in 1974 in Mumbai. She started in Bollywood with director Jaidev, whom she met in Delhi, got her the chance to play at the inauguration of Doordarshan. She first worked with the music composer Kalyanji-Anandji for the title song of a film called Ek Se Badhkar Ek (1976). She gained popularity in India with the songs "O Mera Babu Chail Chabila" and "Dama Dam Mast Qalandar". In 1974, she recorded "Shaadher Lau" in Kolkata. Laila's name has been written on the Guinness World Records for recording 30 songs within 3 days. In 1982, she won Golden Disk Award as her album Superuna composed by Bappi Lahiri was sold over 1 lakh copies on the first day of its release.


title: Runa Laila

text: In October 2009, she released "Kala Sha Kala", a collection of Punjabi wedding songs, in India. In 2012, Laila served as a judge on the show Sur Kshetra, an Indian television contest show for amateur singers. She described her relationship with fellow judge Asha Bhosle as that of sisters. In 2014, she collaborated with Sabina Yasmin on a song for a television play ""Dalchhut Projapoti"", the first time they worked on a song together. Runa has sung in seventeen languages including her native Bengali, Hindi, Urdu, Punjabi, Sindhi, Gujarati, Pashto, Baluchi, Arabic, Persian, Malay, Nepalese, Japanese, Italian, Spanish, French and English.


title: Runa Laila

text: Laila has been married three times. She first married Khawaza Javed Kaiser, secondly a Swiss citizen named Ron Daniel and then actor Alamgir. She has a daughter Tani. Her grandson Zain Islam had been selected for the Arsenal progression center in 2012 when he was eight. Her other grandson Aaron Islam is also there.


title: Runa Laila

text: After her sister died in 1976 from cancer, Laila held several charity concerts in Dhaka. The money raised was used to build a cancer hospital in Dhaka. Laila was named a SAARC Goodwill Ambassador for HIV/AIDS. She is the first Bangladeshi to hold this post. She visited New Delhi in 2013 on her first trip as the SAARC ambassador. She met India's External and Health ministers.


title: Elaeis guineensis

text: Elaeis guineensis is a species of palm commonly just called oil palm but also sometimes African oil palm or macaw-fat. It is the principal source of palm oil. It is native to west and southwest Africa, specifically the area between Angola and the Gambia; the species name, "guineensis", refers to the name for the area, Guinea, and not the modern country now bearing that name. The species is also now naturalised in Madagascar, Sri Lanka, Malaysia, Indonesia, Central America, Cambodia, the West Indies, and several islands in the Indian and Pacific Oceans. The closely related American oil palm "Elaeis oleifera" and a more distantly related palm, "Attalea maripa", are also used to produce palm oil.


title: Elaeis guineensis

text: "E. guineensis" was domesticated in West Africa along the south-facing Atlantic coast. There is insufficient documentation and insufficient research to make any guesses as to when this occurred. Human use of oil palms may date as far back as 5,000 years in Egypt; in the late 1800s, archaeologists discovered palm oil in a tomb at Abydos dating back to 3000 BCE.


title: Elaeis guineensis

text: The first Western person to describe it and bring back seeds was the French naturalist Michel Adanson.


title: Elaeis guineensis

text: Oil palms can produce much more oil per unit of land area than most other oil-producing plants (about nine times more than soy and 4.5 times more than rapeseed).


title: Elaeis guineensis

text: "E. guineensis" is monocotyledonous. Mature palms are single-stemmed and grow to tall. The leaves are pinnate and reach long. A young palm produces about 30 leaves a year. Established palms over 10 years produce about 20 leaves a year. The flowers are produced in dense clusters; each individual flower is small, with three sepals and three petals.


title: Elaeis guineensis

text: The palm fruit takes 5–6 months to develop from pollination to maturity. It is reddish, about the size of a large plum, and grows in large bunches. Each fruit is made up of an oily, fleshy outer layer (the pericarp), with a single seed (the palm kernel), also rich in oil. When ripe, each bunch of fruit weighs between depending on the age of the palm tree.


title: Elaeis guineensis

text: For each hectare of oil palm, which is harvested year-round, the annual production averages 20 tonnes of fruit yielding of palm oil and of seed kernels yielding of high-quality palm kernel oil, as well as of kernel meal. Kernel meal is processed for use as livestock feed.


title: Elaeis guineensis

text: All modern, commercial planting material consists of tenera palms or DxP hybrids, which are obtained by crossing thickshelled dura with shell-less pisifera. Although common commercial germinated seed is as thick-shelled as the dura mother palm, the resulting palm will produce thin-shelled tenera fruit. An alternative to germinated seed, once constraints to mass production are overcome, are tissue-cultured or "clonal" palms, which provide "true copies" of high-yielding DxP palms.


title: Elaeis guineensis

text: The Asian effective population size is very limited. The cultivars comprising cultivation in Asia descend from only four trees, which are themselves probably the result of a selfing of one parent.


title: Elaeis guineensis

text: Several varieties and forms of "Elaeis guineensis" have been selected that have different characteristics. These include:


title: Elaeis guineensis

text: Before the Second World War, selection work had started in the "Deli dura" population in Malaya. Pollen was imported from Africa, and DxT and DxP crosses were made. Segregation of fruit forms in crosses made in the 1950s was often incorrect. In the absence of a good marker gene, there was no way of knowing whether control of pollination was adequate.


title: Elaeis guineensis

text: After the work of Beirnaert and Vanderweyen (1941), it became feasible to monitor the efficacy of controlled pollination. From 1963 until the introduction of the palm-pollinating weevil "Elaeidobius kamerunicus" in 1982, contamination in Malaysia's commercial plantings was generally low. Thrips, the main pollinating agent at that time, apparently rarely gained access to bagged female inflorescences. However, "E. kamerunicus" is much more persistent, and after it was introduced, "Deli dura" contamination became a significant problem. This problem apparently persisted for much of the 1980s, but in a 1991 comparison of seed sources, contamination had been reduced to below 2%, indicating control had been restored.


title: Elaeis guineensis

text: A 1992 study at a trial plot in Banting, Selangor, revealed the "yield of Deli dura oil palms after four generations of selection was 60% greater than that of the unselected base population. Crossing the dura and pisifera to give the thin-shelled tenera fruit type improved partitioning of dry matter within the fruit, giving a 30% increase in oil yield at the expense of shell, without changing total dry matter production."


title: Elaeis guineensis

text: In 2013, the gene responsible for controlling shell thickness was discovered, making it possible to verify tenera (DxP) status while palms are still in the nursery.


title: Elaeis guineensis

text: "E. guineensis" is almost entirely pollinated by insects and not by wind. "Elaeidobius kamerunicus" is the most specially adapted pollination partner in Africa. It has been deliberately introduced into southeast Asia in 1981 and the results have been dramatic – Cik Mohd Rizuan et al., 2013 find good results in in Sabah. Contrary to earlier speculation, the introduced population was not too inbred, and inbreeding depression was not the cause of some incidences of lessened fruit set in SEA. Other causes have been proposed. "E. kamerunicus" and the pollination it provides can be negatively affected by nematodes.


title: Elaeis guineensis

text: Worldwide the two most impactful diseases are "Ganoderma orbiforme" (syn. "Ganoderma boninense", basal stem rot, BSR, reviewed by Chong et al 2017) and "Phytophthora palmivora" (bud rot, reviewed by Torres et al 2016). The earliest stages of data gathering and investigation have been performed for disease resistance breeding however propagation material is not available and full breeding programs are not ongoing .


title: Elaeis guineensis

text: Basal stem rot is the most serious disease of oil palm in Malaysia and Indonesia. Previously, research on basal stem rot was hampered by the failure to artificially infect oil palms with the fungus. Although "Ganoderma" had been associated with BSR, proof of its pathogenicity to satisfy Koch's postulate was only achieved in the early 1990s by inoculating oil palm seedling roots or by using rubber wood blocks. A reliable and quick technique was developed for testing the pathogenicity of the fungus by inoculating oil palm germinated seeds.


title: Elaeis guineensis

text: This fatal disease can lead to losses as much as 80% after repeated planting cycles. "Ganoderma" produces enzymes that degrade the infected xylem, thus causing serious problems to the distribution of water and other nutrients to the top of the palm. "Ganoderma" infection is well defined by its lesion in the stem. The cross-section of infected palm stem shows that the lesion appears as a light brown area of rotting tissue with a distinctive, irregularly shaped, darker band at the borders of this area. The infected tissue become as an ashen-grey powdery and if the palm remains standing, the infected trunk rapidly becomes hollow.


title: Elaeis guineensis

text: In a 2007 study in Portugal, scientists suggested control of the fungus on oil palms would benefit from further consideration of the process as one of white rot. "Ganoderma" is an extraordinary organism capable exclusively of degrading lignin to carbon dioxide and water; celluloses are then available as nutrients for the fungus. It is necessary to consider this mode of attack as a white rot involving lignin biodegradation, for integrated control. The existing literature does not report this area and appears to be concerned particularly with the mode of spread and molecular biology of "Ganoderma". The white rot perception opens up new fields in breeding/selecting for resistant cultivars of oil palms with high lignin content, ensuring the conditions for lignin decomposition are reduced, and simply sealing damaged oil palms to stop decay. The spread likely is by spores rather than roots. The knowledge gained can be employed in the rapid degradation of oil palm waste on the plantation floor by inoculating suitable fungi, and/or treating the waste more appropriately (e.g. chipping and spreading over the floor rather than windrowing).


title: Elaeis guineensis

text: "Phytophthora palmivora" has caused a loss of of "E. guineensis" near San Lorenzo in Ecuador. The protozoa cause bud rot (Spanish: pudrición del cogollo). In reaction, growers there replanted using a hybrid of "E. guineensis" and "E. oleifera", the South American oil palm.


title: Elaeis guineensis

text: Endophytic bacteria are organisms inhabiting plant organs that at some time in their life cycles can colonize the internal plant tissues without causing apparent harm to the host. Introducing endophytic bacteria to the roots to control plant disease is to manipulate the indigenous bacterial communities of the roots in a manner, which leads to enhanced suppression of soil-borne pathogens. The use of endophytic bacteria should thus be preferred to other biological control agents, as they are internal colonizers, with better ability to compete within the vascular systems, limiting "Ganoderma" for both nutrients and space during its proliferation. Two bacterial isolates, "Burkholderia cepacia" (B3) and "Pseudomonas aeruginosa" (P3) were selected for evaluation in the glasshouse for their efficacy in enhancing growth and subsequent suppression of the spread of BSR in oil palm seedlings.


title: Elaeis guineensis

text: Little leaf syndrome has not been fully explained, but has often been confused with boron deficiency. The growing point is damaged, sometimes by "Oryctes" beetles. Small, distorted leaves resembling those due to a boron deficiency emerge. This is often followed by secondary pathogenic infections in the spear that can lead to spear rot and palm death.


title: Elaeis guineensis

text: "M. plana" is a Lepidopteran moth and a major pest of oil palms in Malaysia. "M. plana" outbreaks in Malaysia are highly correlated with relative humidity. Relative humidity estimates based on satellite remote sensing data were fed into both regression models and neural networks. The predictions of both were found to be closely correlated with actual "M. plana" appearance on plantations, with the NN producing the best results.


title: Elaeis guineensis

text: Other arthropods include: Bagworm moths (the Psychidae family), the Asiatic rhinoceros beetle ("Oryctes rhinoceros"), "Rhynchophorus palmarum" (the South American palm weevil), "Tirathaba mundella" (the oil palm bunch moth), and "Tirathaba rufivena" (the coconut spike moth).


title: Elaeis guineensis

text: Besides direct damage to plant material, rats also predate on "Elaeidobius kamerunicus", the African palm pollinating weevil.


title: Elaeis guineensis

text: Chimpanzees ("Pan troglodytes") are known to use stones to crack open the nuts of "E. guineensis". a rare example of tool use by animals.


title: Elaeis guineensis

text: Grey parrots ("Psittacus erithacus") are known to prefer oil palm fruit in the wild. One of their chief predators, the palm-nut vulture ("Gypohierax angolensis"), also heavily depends on oil palm fruit for its diet, making up over 60% of the adult bird's diet and over 90% of the juvenile bird's diet (along with Raffia palm).


title: Elaeis guineensis

text: "Bursaphelenchus cocophilus" is a nematode pest which is better known for infecting coconut palms. (It also afflicts a few other of the Arecaceae.) It causes "red ring disease", so named because it produces a red colored layer within the trunk of the tree, which looks like a red ring in a cross section cut. "B. cocophilus" is obligately transmitted as the third juvenile stage by vectors, specifically several species of weevil. Unlike congener "B. xylophilus" there are "not" thought to be any non plant hosts to serve as reservoir hosts for infection of "E. guineensis". Besides direct infestation of the palm, other nematodes infest the pollinating weevil "Elaeidobius kamerunicus", reducing pollination and yield.


title: Elaeis guineensis

text: Because each tree is relatively big and has an individual value, information about its pest and disease status is valuable. Although visual inspection is the oldest method, others are under development or occasional use.


title: Elaeis guineensis

text: Volatiles and microfocus X-ray fluorescence are two methods can be used to non-invasively detect pre-emergence "Ganoderma orbiforme" disease as a lab test. Sonic tomography is already in use with good results, at 96% accuracy. On the other hand satellite imagery and computer vision has low classification accuracy as to severity.


title: Elaeis guineensis

text: Oil palms were introduced to Java by the Dutch in 1848, and to Malaysia (then the British colony of Malaya) in 1910 by Scotsman William Sime and English banker Henry Darby. The species of palm tree "Elaeis guineensis" was taken to Malaysia from Eastern Nigeria in 1961. As noted it originally grew in West Africa. The southern coast of Nigeria was originally called the Palm oil coast by the first Europeans who arrived there and traded in the commodity. This area was later renamed the Bight of Biafra.


title: Elaeis guineensis

text: In traditional African medicine different parts of the plant are used as laxative and diuretic, as a poison antidote, as a cure for gonorrhea, menorrhagia, and bronchitis, to treat headaches and rheumatism, to promote healing of fresh wounds and treat skin infections.


title: Elaeis guineensis

text: In Cambodia, this palm was introduced as a decorative plant in public gardens, its Khmer name is "dôô:ng préing" (doong=palm, preing=oil).


title: Elaeis guineensis

text: In Malaysia, the first plantations were mostly established and operated by British plantation owners, such as Sime Darby and Boustead, and remained listed in London until the Malaysian government engineered their "Malaysianisation" throughout the 1960s and 1970s.


title: Elaeis guineensis

text: Federal Land Development Authority (Felda) is the world's biggest oil palm planter, with planted area close to 900,000 hectares in Malaysia and Indonesia. Felda was formed on July 1, 1956, when the Land Development Act came into force with the main aim of eradicating poverty. Settlers were each allocated 10 acres of land (about 4 hectares) planted either with oil palm or rubber, and given 20 years to pay off the debt for the land.


title: Elaeis guineensis

text: After Malaysia achieved independence in 1957, the government focused on value-added of rubber planting, boosting exports, and alleviating poverty through land schemes. In the 1960s and 1970s, the government encouraged planting of other crops, to cushion the economy when world prices of tin and rubber plunged. Rubber estates gave way to oil palm plantations. In 1961, Felda's first oil palm settlement opened, with 3.75 km of land. As of 2000, 6855.2 km (approximately 76%) of the land under Felda's programmes were devoted to oil palms. By 2008, Felda's resettlement broadened to 112,635 families, who work on 8533.13 km of agriculture land throughout Malaysia. Oil palm planting took up 84% of Felda's plantation landbank.


title: Elaeis guineensis

text: FELDA's success led to the establishment of other development schemes to support the establishment of small-farmer oil palm cultivation. The Federal Land Consolidation and Rehabilitation Authority (FELCRA) was established in 1966 and the Sarawak Land Consolidation and Rehabilitation Authority (SALCRA) was formed in 1976. The primary objective of these organizations is to assist in the development of rural communities and reduce poverty through the cultivation of high yielding crops such as palm oil.


title: Elaeis guineensis

text: , SALCRA had developed 18 estates totalling approximately 51,000 hectares. That year the organization shared dividends with 16,374 landowners participating in the program.


title: Elaeis guineensis

text: Oil is extracted from both the pulp of the fruit (palm oil, an edible oil) and the kernel (palm kernel oil, used in foods and for soap manufacture). For every 100 kg of fruit bunches, typically 22 kg of palm oil and 1.6 kg of palm kernel oil can be extracted.


title: Elaeis guineensis

text: The high oil yield of oil palms (as high as 7,250 liters per hectare per year) has made it a common cooking ingredient in Southeast Asia and the tropical belt of Africa. Its increasing use in the commercial food industry in other parts of the world is buoyed by its cheaper pricing, the high oxidative stability of the refined product, and high levels of natural antioxidants.


title: Elaeis guineensis

text: The oil palm originated in West Africa, but has since been planted successfully in tropical regions within 20 degrees of the equator. In the Republic of the Congo, or Congo Brazzaville, precisely in the Northern part, not far from Ouesso, local people produce this oil by hand. They harvest the fruit, boil it to let the water evaporate, then press what is left to collect the reddish-orange-colored oil.


title: Elaeis guineensis

text: In 1995, Malaysia was the world's largest producer, with a 51% of world share, but since 2007, Indonesia has been the world's largest producer, supplying approximately 50% of world palm oil volume.


title: Elaeis guineensis

text: In 2019 total production was "E. guineensis" is among the few tropical tree crops (along with bananas and citrus) with high productivity in actual growing conditions, i.e. outside of test plots.


title: Elaeis guineensis

text: The social and environmental impact of oil palm cultivation is a highly controversial topic. Oil palm is a valuable economic crop and provides a major source of employment. It allows many small landholders to participate in the cash economy and often results in the upgrade of the infrastructure (schools, roads, telecommunications) within that area. According to the IBGE oil palm is a common crop in agroforestry practices in the Amazon. However, there are cases where native customary lands have been appropriated by oil palm plantations without any form of consultation or compensation, leading to social conflict between the plantations and local residents. In some cases, oil palm plantations are dependent on imported labour or illegal immigrants, with some concerns about the employment conditions and social impacts of these practices.


title: Elaeis guineensis

text: Biodiversity loss (including the potential extinction of charismatic species) is one of the most serious negative effects of oil palm cultivation. On the other hand, it also helps to push invasive species further, e.g. "Anoplolepis gracilipes" in southeast Asia. Large areas of already threatened tropical rainforest are often cleared to make way for palm oil plantations, especially in Southeast Asia, where enforcement of forest protection laws is lacking. In some states where oil palm is established, lax enforcement of environmental legislation leads to encroachment of plantations into protected areas, encroachment into riparian strips, open burning of plantation wastes, and release of palm mill pollutants such as palm oil mill effluent (POME) in the environment. Some of these states have recognised the need for increased environmental protection, resulting in more environment-friendly practices. Among those approaches is anaerobic treatment of POME, which can be a good source for biogas (methane) production and electricity generation. Anaerobic treatment of POME has been practiced in Malaysia and Indonesia. Like most wastewater sludge, anaerobic treatment of POME results in dominance of "Methanosaeta concilii". It plays an important role in methane production from acetate, and the optimum condition for its growth should be considered to harvest biogas as renewable fuel.


title: Elaeis guineensis

text: Demand for palm oil has increased in recent years due to its use as a biofuel, but recognition that this increases the environmental impact of cultivation, as well as causing a food vs fuel issue, has forced some developed nations to reconsider their policies on biofuel to improve standards and ensure sustainability. However, critics point out that even companies signed up to the Roundtable on Sustainable Palm Oil continue to engage in environmentally damaging practices and that using palm oil as biofuel is perverse because it encourages the conversion of natural habitats such as forests and peatlands, releasing large quantities of greenhouse gases.


title: Elaeis guineensis

text: Oil palm production has been documented as a cause of substantial and often irreversible damage to the natural environment. Its impacts include deforestation, habitat loss of critically endangered species, and a significant increase in greenhouse gas emissions.


title: Elaeis guineensis

text: The pollution is exacerbated because many rainforests in Indonesia and Malaysia lie atop peat bogs that store great quantities of carbon, which are released when the forests are cut down and the bogs are drained to make way for the plantations.


title: Elaeis guineensis

text: Environmental groups, such as Greenpeace, claim the deforestation caused by making way for oil palm plantations is far more damaging for the climate than the benefits gained by switching to biofuel. Fresh land clearances, especially in Borneo, are contentious for their environmental impact.


title: Elaeis guineensis

text: Despite thousands of square kilometres of land standing unplanted in Indonesia, tropical hardwood forests are being cleared for palm oil plantations. Furthermore, as the remaining unprotected lowland forest dwindles, developers are looking to plant peat swamp land, using drainage that begins an oxidation process of the peat which can release 5,000 to 10,000 years worth of stored carbon. Drained peat is also at very high risk of forest fire. There is a clear record of fire being used to clear vegetation for oil palm development in Indonesia, where in recent years drought and man-made clearances have led to massive uncontrolled forest fires, covering parts of Southeast Asia in haze and leading to an international crisis with Malaysia. These fires have been blamed on a government with little ability to enforce its own laws, while impoverished small farmers and large plantation owners illegally burn and clear forests and peat lands to develop the land rather than reap the environmental benefits it could offer.


title: Elaeis guineensis

text: Many of the major companies in the vegetable oil economy participate in the Roundtable on Sustainable Palm Oil, which is trying to address this problem. For example, in 2008, Unilever, a member of the group, committed to use only oil palm oil which is certified as sustainable, by ensuring the large companies and smallholders that supply it convert to sustainable production by 2015.


title: Elaeis guineensis

text: Meanwhile, much of the recent investment in new palm plantations for biofuel has been funded through carbon credit projects through the Clean Development Mechanism; however, the reputational risk associated with the unsustainable palm plantations in Indonesia has now made many funds wary of such investment.


title: Elaeis guineensis

text: Some scientists and companies are going beyond using just the oil, and are proposing to convert fronds, empty fruit bunches and palm kernel shells harvested from oil palm plantations into renewable electricity, cellulosic ethanol, biogas, biohydrogen and bioplastic. Thus, by using both the biomass from the plantation as well as the processing residues from palm oil production (fibers, kernel shells, palm oil mill effluent), bioenergy from palm plantations can have an effect on reducing greenhouse gas emissions. Examples of these production techniques have been registered as projects under the Kyoto Protocol's Clean Development Mechanism.


title: Elaeis guineensis

text: By using palm biomass to generate renewable energy, fuels and biodegradable products, both the energy balance and the greenhouse gas emissions balance for palm biodiesel is improved. For every tonne of palm oil produced from fresh fruit bunches, a farmer harvests around 6 tonnes of waste palm fronds, 1 tonne of palm trunks, 5 tonnes of empty fruit bunches, 1 tonne of press fiber (from the mesocarp of the fruit), half a tonne of palm kernel endocarp, 250 kg of palm kernel press cake, and 100 tonnes of palm oil mill effluent. Some oil palm plantations incinerate biomass to generate power for palm oil mills. Some other oil palm plantations yield large amount of biomass that can be recycled into medium density fibreboards and light furniture. In efforts to reduce greenhouse gas emissions, scientists treat palm oil mill effluent to extract biogas. After purification, biogas can substitute for natural gas for use at factories. Anaerobic treatment of palm oil mill effluent, practiced in Malaysia and Indonesia, results in domination of "Methanosaeta concilii". It plays an important role in methane production from acetate and the optimum condition for its growth should be considered to harvest biogas as renewable fuel.


title: Elaeis guineensis

text: Unfortunately, the production of palm oil has detrimental effects on the environment and is not considered to be a sustainable biofuel. The deforestation occurring throughout Malaysia and Indonesia as a result of the growing demand for this plant has made scarce natural habitats for orangutans and other rainforest dwellers. More carbon is released during the life cycle of a palm oil plant to its use as a biofuel than is emitted by the same volume of fossil fuels.


title: Double First Class University Plan

text: The World First Class University and First Class Academic Discipline Construction (), together known as Double First Class (), is a tertiary education development initiative designed by the People's Republic of China central government in 2015, which aims to comprehensively develop elite Chinese universities into world-class institutions by the end of 2050 through developing and strengthening their individual faculty departments. The Double First-Class University Plan has made new arrangements for the Chinese higher education institution development. The universities included in this plan are called Double First Class Universities.


title: Double First Class University Plan

text: The Double First-Class Universities are considered to be the most elite institutions of Chinese tertiary education, representing the top 5% of overall universities and colleges in Mainland China (approximately 3,000 higher education institutions).


title: Double First Class University Plan

text: In October 2015, the State Council of China published the 'Overall Plan for Promoting the Construction of World First Class Universities and First Class Disciplines' (Double First Class University Plan), which made new arrangements for the development of higher education in China, replacing previous higher education projects.


title: Double First Class University Plan

text: In June 2016, the Ministry of Education of China announced that the Project 211 and Project 985 had been abolished and replaced by the Double First Class University Initiative.


title: Double First Class University Plan

text: In September 2017, the full list of the universities and their disciplines of the Double First Class University Plan was jointly published by the Ministry of Education of China, the Ministry of Finance of China and the National Development and Reform Commission of China. According to the list, 140 universities have been approved as ‘Double First Class Universities’ by the State Council of China. The Double First Class University Plan underlines that driving the overall development of universities by building and strengthening their faculties and departments, and eventually developing the 140 listed elite universities into world-class universities by 2050.


title: Double First Class University Plan

text: The Double First Class University Plan represents a new way of ranking universities in China. By 2021, the 140 Double First Class Universities accounted for roughly 4.65% of the 3,012 universities and colleges in Mainland China, representing the most elite part of higher education institutions in this country.


title: Double First Class University Plan

text: In December 2021, the 23rd meeting of Central Comprehensively Deepening Reforms Commission chaired by General Secretary of the Chinese Communist Party Xi Jinping deliberated and approved 'Some Suggestions on Further Promoting the Construction of World First Class Universities and First Class Disciplines'.


title: Double First Class University Plan

text: In February 2022, the 'Some Suggestions on Further Promoting the Construction of World First Class Universities and First Class Disciplines' were jointly published by the Ministry of Education, the Ministry of Finance, and the National Development and Reform Commission. In addition, with the approval of the State Council of China, the updated list of "Double First Class Universities" has been released. According to the Phase 2 List of the Double First Class University Plan, the previous university classifications have been abolished, and a total number of 147 universities have been included in the plan (accounted for 4.88% of 3,012 Chinese higher education institutions), after 7 universities newly adding into the list since February 2022.


title: Double First Class University Plan

text: The Phase 1 list of 140 Double First Class Universities were issued by Chinese Ministry of Education, Ministry of Finance, and National Development and Reform Commission without any ranking within the lists (by "school code") in 2017. In February 2022, the Ministry of Education of China announced that the classifications had been invalidated, and the universities included in the Double First Class University Plan would be referred to as Double First Class Universities. According to the updated Phase 2 list of the Double First Class University Plan (without any ranking within the list), a total of 147 universities have been approved and recognized as Double First Class Universities, taking up 4.88% of universities and colleges in China.


title: Double First Class University Plan

text: <nowiki>*</nowiki> represent the newly added universities in the Phase 2 list of Double First Class University Plan.


title: Double First Class University Plan

text: The Double First Class University Plan underlines that developing the listed Chinese elite universities into world-class universities by 2050 through strengthening their developments of disciplines and faculties.


title: Double First Class University Plan

text: As of 2022,  110 Double First-Class universities among the Chinese mainland universities shortlisted in the world's top 1,000 by the Academic Ranking of World Universities, including 61 Top 500 universities, 38 Top 300 universities, 26 Top 200 universities and 8 Top 100 universities.


title: Double First Class University Plan

text: On 18 September 2020, the members of "Double First Class" expert valuation group, which was headed by Lin Huiqing, Chairman of the Medical Education Expert Committee of the Ministry of Education and former Vice Minister of the Ministry of Education, unanimously agreed that Tsinghua University has been fully established as a world-class university.


title: Evolution of the wolf

text: The grey wolf "Canis lupus" is a highly adaptable species that is able to exist in a range of environments and which possesses a wide distribution across the Holarctic. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other. This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.


title: Evolution of the wolf

text: The archaeological and paleontological records show grey wolf continuous presence for at least the last 300,000 years. This continuous presence contrasts with genomic analyses, which suggest that all modern wolves and dogs descend from a common ancestral wolf population that existed as recently as 20,000 years ago. These analyses indicate a population bottleneck, followed by a rapid radiation from an ancestral population at a time during, or just after, the Last Glacial Maximum. This implies that the original wolf populations were out-competed by a new type of wolf which replaced them. However, the geographic origin of this radiation is not known.


title: Evolution of the wolf

text: The fossil record for ancient vertebrates is composed of rarely occurring fragments from which it is often impossible to obtain genetic material. Researchers are limited to morphologic analysis but it is difficult to estimate the intra-species and inter-species variations and relationships that existed between specimens across time and place. Some observations are debated by researchers who do not always agree, and hypotheses that are supported by some authors are challenged by others.


title: Evolution of the wolf

text: The Cretaceous–Paleogene extinction event occurred 65 million years ago and brought an end to the dinosaurs and the appearance of the first carnivorans. The name carnivoran is given to a member of the order Carnivora. Carnivorans possess a common arrangement of teeth called carnassials, in which the first lower molar and the last upper premolar possess blade-like enamel crowns that act similar to a pair of shears for cutting meat. This dental arrangement has been modified by adaptation over the past 60 million years for diets composed of meat, for crushing vegetation, or for the loss of the carnassial function altogether as in seals, sea lions, and walruses. Today, not all carnivorans are carnivores, such as the insect-eating Aardwolf.


title: Evolution of the wolf

text: The carnivoran ancestors of the dog-like caniforms and the cat-like feliforms began their separate evolutionary paths just after the end of the dinosaurs. The first members of the dog family Canidae appeared 40 million years ago, of which only its subfamily the Caninae survives today in the form of the wolf-like and fox-like canines. The caniforms included the fox-like genus "Leptocyon" whose various species existed from 34 million YBP before branching 11.9 million YBP into "Vulpes" (foxes) and Canini (canines). The jackal-sized "Eucyon" existed in North America from 10 million YBP and by the Early Pliocene about 6–5 million YBP the coyote-like "Eucyon davisi" invaded Eurasia. In North America it gave rise to early "Canis" which first appeared in the Miocene (6 million YBP) in south-western US and Mexico. By 5 million YBP the larger "Canis lepophagus" appeared in the same region.


title: Evolution of the wolf

text: The canids that had immigrated from North America to Eurasia – "Eucyon", "Vulpes", and "Nyctereutes" – were small to medium-sized predators during the Late Miocene and Early Pliocene but they were not the top predators. The position of the canids would change with the arrival of "Canis" to become a dominant predator across the Holarctic. The wolf-sized "C. chihliensis" appeared in northern China in the Mid-Pliocene around 4–3 million YBP. The large wolf-sized "Canis" appeared in the Middle Pliocene about 3 million years ago in the Yushe Basin, Shanxi Province, China. By 2.5 million years ago its range included the Nihewan Basin in Yangyuan County, Hebei, China and Kuruksay, Tadzhikistan. This was followed by an explosion of "Canis" evolution across Eurasia in the Early Pleistocene around 1.8 million YBP in what is commonly referred to as the "wolf event". It is associated with the formation of the mammoth steppe and continental glaciation. "Canis" spread to Europe in the forms of "C. arnensis", "C. etruscus", and "C. falconeri". Other studies state that the oldest "Canis" remains that have been found in Europe were from France and dated to 3.1 million YBP, followed by "Canis cf. etruscus" (where cf. in Latin means confer, uncertain) from Italy dated to 2.2 million YBP.


title: Evolution of the wolf

text: The fossil record is incomplete but it is likely that wolves arose from a population of small, early canids. Morphological evidence and genetic evidence both suggest that wolves evolved during the Pliocene and Early Pleistocene eras from the same lineage that also led to the coyote, with fossil specimens indicating that the coyote and the wolf diverged from a common ancestor 1.5 million years ago. The ancestor of the jackal and the other extant members of the genus "Canis" had split from the lineage before this time.


title: Evolution of the wolf

text: After this separation from a common ancestor the species that were believed to be involved in the further evolution of the wolf and coyote – and the beliefs of some paleontologists – diverged. A number of researchers believed that the lines of "C. priscolatrans", "C. etruscus", "C. rufus", "C. lycaon", and "C. lupus" were components involved in some way that lead to the modern wolf and coyote.


title: Evolution of the wolf

text: "Canis lepophagus" lived in the early Pliocene in North America. Kurten proposed that the Blancan "C. lepophagus" derived from smaller Miocene "Canis" species in North America. It then became widespread across Eurasia where it was either identical to, or closely related with, "C. arnensis" of Europe.


title: Evolution of the wolf

text: Johnston describes "C. lepophagus" as having a more slender skull and skeleton than in the modern coyote. Robert M. Nowak found that the early populations had small, delicate and narrowly proportioned skulls that resemble small coyotes and appear to be ancestral to "C. latrans". Johnson noted that some specimens found in Cita Canyon, Texas, had larger, broader skulls, and along with other fragments Nowak suggested that these were evolving into wolves.


title: Evolution of the wolf

text: Tedford disagreed with previous authors and found that its cranio-dental morphology lacked some characteristics that are shared by "C. lupus" and "C. latrans", and therefore there was not a close relationship but it did suggest "C. lepophagus" was the ancestor of both wolves and coyotes.


title: Evolution of the wolf

text: "Canis priscolatrans" lived in the late Pliocene-Early Pleistocene in North America. The first definite wolf appeared in the Late Blancan/Early Irvingtonian, and named "C. priscolatrans" that was either very close to or a synonym for "Canis edwardii". It resembled "C. rufus" in cranial size and proportions but with more complex dentition. However, there are no fossils of "C. rufus" until the Late Rancholabrean.


title: Evolution of the wolf

text: Kurtén was uncertain if "C. priscolatrans" derived from "C. lepophagus" and "C. arnensis", but believed that "C. priscolatrans" was a population of large coyotes that were ancestral to Rancholabrean and recent "C. latrans". He noted that "C. arnensis" of Europe showed striking similarities to "C. priscolatrans", and they could represent what once was a holarctic population of coyotes. Nowak disagreed, and believed that "C. priscolatrans" was a counterpart to the European "C. etruscus". Kurtén later proposed that both "C. priscolatrans" and "C. etruscus" were part of a group which led to "C. lupus" but was not sure if they evolved separately from "C. lepophagus" or a possible common ancestor that was derived from "C. lepophagus".


title: Evolution of the wolf

text: The remains of the larger coyote-like "Canis edwardii" have been found in the later Pliocene in the south-western USA along with "C. lepophagus", which indicates a descent. Tedford recognised "C. edwardii" and found that the cranio-dental morphology of "C. priscolatrans" fell inside that of "C. edwardii" such that the species name "C. priscolatrans" was doubtful ("nomen dubium").


title: Evolution of the wolf

text: The North American wolves became larger, with tooth specimens indicating that "C. priscolatrans" diverged into the large wolf "C. armbrusteri". during the Middle Pleistocene in North America. Robert A. Martin disagreed, and believed that "C. armbrusteri" was "C. lupus". Nowak disagreed with Martin and proposed that "C. armbrusteri" was not related to "C. lupus" but "C. priscolatrans", which then gave rise to "A. dirus". Tedford proposed that the South American "C. gezi" and "C. nehringi" share dental and cranial similarities developed for hypercarnivory, suggesting "C. armbrusteri" was the common ancestor of "C. gezi", "C. nehringi" and "A. dirus".


title: Evolution of the wolf

text: In 1908 the paleontologist John Campbell Merriam began retrieving numerous fossilized bone fragments of a large wolf from the Rancho LaBrea tar pits. By 1912 he had found a skeleton sufficiently complete to be able to formally recognize these and the previously found specimens under the name "C.dirus" (Leidy 1858).


title: Evolution of the wolf

text: "Canis dirus", or as it is now widely recognized, "Aenocyon dirus", lived in the late Pleistocene to early Holocene in North and South America. In 1987, a new hypothesis proposed that a mammal population could give rise to a larger form called a hypermorph during times when food was abundant, but when food later became scarce the hypermorph would either adapt to a smaller form or go extinct. This hypothesis might explain the large body sizes found in many Late Pleistocene mammals compared to their modern counterparts. Both extinction and speciation – a new species splitting from an older one – could occur together during periods of climatic extremes. GloriaD.Goulet agreed with Martin and further proposed that this hypothesis might explain the sudden appearance of "A.dirus" in North America, and that because of the similarities in their skull shapes that "C.lupus" gave rise to the "A.dirus" hypermorph due to abundant game, a stable environment, and large competitors. Nowak, Kurten and Berta disagreed with Goulet and proposed that "A. dirus" was not derived from "C. lupus". The three noted paleontologists Xiaoming Wang, R. H. Tedford and R. M. Nowak have all proposed that "A. dirus" had evolved from "C. armbrusteri", with Nowak stating that there were specimens from Cumberland Cave, Maryland, that indicated "C. armbrusteri" diverging into "A. dirus". The two taxa share a number of characteristics (synapomorphy), which suggests an origin of "A. dirus" in the late Irvingtonian in the open terrain in the midcontinent, and then later expanding eastward and displacing its ancestor "C. armbrusteri".


title: Evolution of the wolf

text: However, in 2021, a study indicated the dire wolf to be a highly divergent lineage which last shared a most recent common ancestor with the wolf-like canines 5.7 million years ago. The morphological similarity between dire wolves and gray wolves was concluded to be due to convergent evolution. This finding indicates that the wolf and coyote lineages evolved in isolation from the dire wolf lineage. The study proposes an early origin of the dire wolf lineage in the Americas, and that this geographic isolation allowed them to develop a degree of reproductive isolation since their divergence 5.7 million years ago. Coyotes, dholes, gray wolves, and the extinct "Xenocyon" evolved in Eurasia and expanded into North America relatively recently during the Late Pleistocene, therefore there was no admixture with the dire wolf. As a result, the study found that the correct binomial name of the dire wolf is "Aenocyon dirus", as proposed by Merriam in 1918. The long-term isolation of the dire wolf lineage implies that other American fossil taxa, including "C. armbrusteri" and "C. edwardii", may also belong to the dire wolf's lineage.


title: Evolution of the wolf

text: Merriam named 3 unusual species based on specimens recovered from the Rancho La Brea tar pits. They were regarded by Nowak as taxonomic synonyms for "Canis lupus".


title: Evolution of the wolf

text: "Canis occidentalis furlongi" (Merriam 1910) was described as a wolf considerably smaller than the dire wolf and more closely related to the timber wolf "Canis lupus occidentalis". However, its premolar P4 (upper carnassials) were massive, and the hypocone of the molar M1 was larger than that of the dire wolf. One specimen's teeth and palate are described as being between the dire wolf and "Canis lupus occidentalis". Nowak proposed the name "Canis lupus furlongi" as he believed that it was a subspecies of the grey wolf. There is the possibility that wolves living in marginal areas led to dire wolf/grey wolf hybrids.


title: Evolution of the wolf

text: "Canis milleri" (Merriam 1912), the Miller wolf, was as large as the timber wolf but with a shorter and heavier head. Its skull and dentition were described as being intermediate between "Canis lupus occidentalis" and the dire wolf. Its skull differed from "occidentalis" due to its wider skull, especially at the palate, and the size of its P4 and M1 were much larger than any known timber wolf, with the P4 approaching that of the dire wolf in size. It is regarded by Nowak as a taxonomic synonym of "Canis lupus furlongi".


title: Evolution of the wolf

text: "Aenocyon milleri" (Merriam 1918) was described as a wolf different from the dire wolf by its smaller size, low sagittal crest, and a less prominent inion, but closer to the dire wolf than the timber wolf. Only one specimen was found. It is regarded by Nowak as a taxonomic synonym of "Canis lupus furlongi".


title: Evolution of the wolf

text: "Canis mosbachensis", sometimes known as the Mosbach wolf, is an extinct small wolf that once inhabited Eurasia from the Middle to Late Pleistocene. The phylogenetic descent of the extant wolf "C. lupus" from "C. etruscus" through "C. mosbachensis" is widely accepted. In 2010, a study found that the diversity of the "Canis" group decreased by the end of the Early Pleistocene to Middle Pleistocene and was limited in Eurasia to the small wolves of the "C. mosbachensis–C. variabilis" group that were a comparable size to the extant Indian wolf ("Canis lupus pallipes"), and the large hypercarnivorous "Canis (Xenocyon) lycaonoides" that was comparable in size to extant northern grey wolves.


title: Evolution of the wolf

text: In 2012, a study of the wolf-like "Canis" species of ancient China under the direction of Xiaoming Wang found that these were all quite close to "C. lupus" in both dental and post-cranial dimensions except for "Canis variabilis", which was "very strange" compared to other "Canis" in China as it had much smaller cranio-dental dimensions than earlier and later species. The study concluded that "It is very likely that this species is the ancestor of the domestic dog "Canis familiaris", a hypothesis that has been proposed by previous authors."


title: Evolution of the wolf

text: Wang and Tedford proposed that the genus "Canis" was the descendant of the coyote-like "Eucyon davisi", and its remains first appeared in the Miocene (6 million YBP) in south-western USA and Mexico. By the Pliocene (5 million YBP), the larger "Canis lepophagus" appeared in the same region and by the Early Pleistocene (1 million YBP) "Canis latrans" (the coyote) was in existence. They proposed that the progression from "Eucyon davisi" to "C lepophagus" to the coyote was linear evolution. Additionally, "C. edwardii", "C. latrans" and "C. aureus" form together a small clade and because "C. edwardii" appeared earliest spanning the mid-Blancan (late Pliocene) to the close of the Irvingtonian (late Pleistocene) it is proposed as the ancestor.


title: Evolution of the wolf

text: Nowak and Tedford also believed that it was possible for "C. lupus" to have been derived from a Miocene or Pliocene canid line that preceded and was separate from "C. lepophagus". Based on morphology from China, the Pliocene wolf "C. chihliensis" may have been the ancestor for both "C. armbrusteri" and "C. lupus" before their migration into North America. "C. chihliensis" appears to be more primitive and smaller than "C. lupus", and measurements of its skull and teeth are similar to "C. lupus" but those of its postcranial elements are smaller. "C. armbrusteri" appeared in North America in the Middle Pleistocene and is a wolf-like form larger than any "Canis" at that time. At the end of the most recent glacial retreat during the past 30,000 years, warming melted the glacial barriers across northern Canada allowing arctic mammals to extend their range into mid-latitude North America, including elk, caribou, bison, and the grey wolf.


title: Evolution of the wolf

text: In Eurasia during the Middle Pleistocene, "C. falconeri" gave rise to the hypercarnivore genus "Xenocyon", which then gave rise to genus "Cuon" (the dhole) and genus "Lycaon" (the African hunting dog). Just before the appearance of "C. dirus", North America was invaded by genus "Xenocyon" that was as large as "C. dirus" and more hypercarnivorous. The fossil record shows them as rare and it is assumed that they could not compete with the newly derived "C. dirus".


title: Evolution of the wolf

text: The large wolf "C. antonii" from late Pliocene to early Pleistocene China was assessed as being a variation within "C. chihliensis", and the large wolf "C. falconeri" occurred abruptly in Europe in the Early Pleistocene, perhaps representing a westward extension of "C. antonii".


title: Evolution of the wolf

text: The earliest "Canis lupus" specimen was a fossil tooth discovered at Old Crow, Yukon, Canada. The specimen was found in sediment dated 1 million YBP, however the geological attribution of this sediment is questioned. Slightly younger specimens were discovered at Cripple Creek Sump, Fairbanks, Alaska, in strata dated 810,000 YBP. Both discoveries point to an origin of these wolves in east Beringia during the Middle Pleistocene.


title: Evolution of the wolf

text: In France, the subspecies "C. l. lunellensis" Bonifay, 1971 discovered at Lunel-Viel, Hérault dated 400–350,000 YBP, "C. l. santenaisiensis" Argant, 1991 from Santenay, Côte-d'Or dated to 200,000 YBP, and "C. lupus maximus" Boudadi-Maligne, 2012 from Jaurens cave, Nespouls, Corrèze dated 31,000 YBP, show a progressive increase in size and are proposed to be chrono-subspecies. In Italy, the earliest "Canis lupus" specimens were found at La Polledrara di Cecanibbio, 20 km north-west of Rome in strata dated 340,000–320,000 YBP. In 2017, a study found that the dimensions of the upper and lower carnassial teeth of the early Holocene Italian wolf are close to those of "C. l. maximus". Fluctuations in the size of "C. lupus" carnassial teeth correlate with the spread of megafauna. The Italian wolf underwent a reduction in body size with the loss of the red deer in Italy during the Renaissance. The proposed lineage is:


title: Evolution of the wolf

text: In 2020, a new species of wolf "C. borjgali" was discovered in Georgia. This wolf is proposed to be the next evolutionary step after "C. mosbachensis" and an ancestor of the "C. lupus" lineage.


title: Evolution of the wolf

text: There are a number of recently discovered specimens which are proposed as being Paleolithic dogs, however their taxonomy is debated. These have been found in either Europe or Siberia and date 40,000-17,000 YBP. They include Hohle Fels in Germany, Goyet Caves in Belgium, Predmosti in the Czech Republic, and four sites in Russia: Razboinichya Cave, Kostyonki-8, Ulakhan Sular, and Eliseevichi 1. Paw-prints from Chauvet Cave in France dated 26,000 YBP are suggested as being those of a dog, however these have been challenged as being left by a wolf. Paleolithic dogs were directly associated with human hunting camps in Europe over 30,000 (YBP) and it is proposed that they were domesticated. They are also proposed to be either a proto-dog and the ancestor of the domestic dog or a type of wolf unknown to science.


title: Evolution of the wolf

text: In 2002, a study was undertaken into the fossil skulls of two large canids that had been found buried within meters of the doorway of what was once a mammoth-bone hut at the Eliseevichi-I Upper Paleolithic site in the Bryansk Region on the Russian Plain, and using an accepted morphologically based definition of domestication declared them to be "Ice Age dogs". The carbon dating gave a calendar-year age estimate that ranged between 16,945–13,905 YBP. In 2013, a study looked at one of these skulls and its mitochondrial DNA sequence was identified as "Canis familiaris".


title: Evolution of the wolf

text: In 2015, a zooarchaeologist stated that "In terms of phenotypes, dogs and wolves are fundamentally different animals."


title: Evolution of the wolf

text: In 1986, a study of skull morphology found that the domestic dog is morphologically distinct from all other canids except the wolf-like canids. "The difference in size and proportion between some breeds are as great as those between any wild genera, but all dogs are clearly members of the same species." In 2010, a study of dog skull shape compared to extant carnivorans proposed that "The greatest shape distances between dog breeds clearly surpass the maximum divergence between species in the Carnivora. Moreover, domestic dogs occupy a range of novel shapes outside the domain of wild carnivorans."


title: Evolution of the wolf

text: The domestic dog compared to the modern wolf shows the greatest variation in the size and shape of the skull (Evans 1979) that range from 7 to 28 cm in length (McGreevy 2004). Wolves are dolichocephalic (long skulled) but not as extreme as some breeds of such as greyhounds and Russian wolfhounds (McGreevy 2004). Canine brachycephaly (short-skulledness) is found only in domestic dogs and is related to paedomorphosis (Goodwin 1997). Puppies are born with short snouts, with the longer skull of dolichocephalic dogs emerging in later development (Coppinger 1995). Other differences in head shape between brachycephalic and dolichocephalic dogs include changes in the craniofacial angle (angle between the basilar axis and hard palate) (Regodón 1993), morphology of the temporomandibular joint (Dickie 2001), and radiographic anatomy of the cribriform plate (Schwarz 2000).


title: Evolution of the wolf

text: Nowak indicated that orbital angle of the eye socket is an important characteristic defining the difference between the dog and the wolf, with the wolf having the lower angle. Nowak compared the orbital angles of four North American canines (including the Indian dog) and produced the following values in degrees: coyote-42.8, wolf-42.8, dog-52.9 dire wolf-53.1. The orbital angle of the eye socket was clearly larger in the dog than in the coyote and the wolf; why it was almost the same as that of the dire wolf was not commented on.


title: Evolution of the wolf

text: Many authors have concluded that compared to the adult extant wolf, the adult domestic dog has a relatively reduced rostrum (front part of the skull), an elevated frontal bone, a wider palate, a broader cranium, and smaller teeth (Hildebrand1954; Clutton-Brock, Corbet & Hills 1976; Olsen 1985; Wayne 1986; Hemmer 1990; Morey 1990). Other authors have disagreed and have stated that these traits can overlap and vary within the two (Crockford 1999; Harrison 1973). Wolf cubs have similar relative skull proportions as adult dogs and this was proposed as evidence that the domestic dog is a neotenic wolf. This was proposed to be due to either human selection for juvenile appearance or due to a pleiotropic effect as a result of selection for juvenile behavior (Clutton-Brock 1977; Belyaev 1979; Wayne 1986; Coppinger and Schneider 1995). Wayne (1986) concluded that his dog samples did not have significant relative shortening of the rostrum compared to wolves, calling this identification feature into question. A 2004 study that used 310 wolf skulls and over 700 dog skulls representing 100 breeds concluded that the evolution of dog skulls can generally not be described by heterochronic processes such as neoteny although some pedomorphic dog breeds have skulls that resemble the skulls of juvenile wolves. "Dogs are not paedomorphic wolves."


title: Evolution of the wolf

text: Compared to the wolf, dog dentition is relatively less robust (Olsen 1985; Hemmer 1990), which is proposed to be due to the relaxation of natural selection when wolves became commensal scavengers, or to artificial selection (Olsen 1985; Clutton-Brock 1995). However, Kieser and Groeneveld (1992) compared the mandibulo-dental measurements of jackals ("C. adustus, C. mesomelas") and Cape foxes ("Vulpes chama") to equivalent-sized dogs and found that the canines of these other canids tended to be slightly smaller and their second molars larger compared to dogs, otherwise the proportions were essentially the same in all species. They concluded that "...the teeth of canids appear to have evolved in concert with one another and relatively independently of differences in dimorphism, size or functional demands". This calls into question the assumption that dog teeth are relatively small due to recent selection, suggesting that dog dentition is plesiomorphic from an ancestor that was smaller than the wolf.


title: Evolution of the wolf

text: The reduced body size of the early dog compared to a wolf is thought due to niche selection (Olsen 1985; Morey 1992; Coppinger &


title: Evolution of the wolf

text: Coppinger 2001). Morey (1992:199) states that "Results...are consistent with a hypothesis that early domestic dogs are evolutionary paedomorphs, products of strong selection for ontogenetically channeled size reduction and alterations of reproductive timing associated with the new domestic way of life." However, in an domestication experiment the domesticated foxes remained the same size as unselected foxes (Trutt 1999:167).


title: Evolution of the wolf

text: Wayne (1986) concluded that the dog is closer in skull morphology to "C. latrans, C. aureus, C. adustus, C. mesomelas, Cuon alpinus" and "Lycaon pictus" than to the wolf. Dahr (1942) concluded that the shape of the dog brain case is closer to that of the coyote than to that of the wolf. Manwell and Baker (1983) reviewed Dahr's work with the addition of dental data for canids and concluded that the dog ancestor was probably within the range of 13.6–20.5 kg, which is smaller than the range 27–54 kg for extant wolves (Mech 1970) and is comparable with the Dingo.


title: Evolution of the wolf

text: The auditory bulla of the dog is relatively smaller and flatter than that of the wolf (Harrison 1973; Clutton-Brock, Corbet & Hill 1976; Nowak 1979; Olsen 1985; Wayne 1986), which is proposed to be due to relaxed selection under domestication as the dog no longer required the acute hearing of the wolf. However, bulla shape has been shown to facilitate increased sensitivity to specific frequencies but shape and size may not be correlated with acuity (Ewer 1973). Therefore, the observed difference could be that the dog bulla has retained its ancestral shape.


title: Evolution of the wolf

text: The ventral edge of the dog's horizontal ramus of the mandible has a convex curve that does not exist in the wolf (Olsen 1985; Clutton-Brock 1995), and no discussion of this difference could be found in the literature. However, Biknevicius and Van Valkenburgh (1997) noticed that the horizontal ramus of bone-processing predators is thicker dorso-ventrally at the point caudal to the site of bone processing. This thickening may have been a function for niche adaptation by the dog's ancestor.


title: Evolution of the wolf

text: A description of the superficial brain morphology of jackals ("C. mesomelas, C. aureus"), coyotes ("C. latrans"), wolves ("C. lupus, C. rufus"), and dogs indicated that the cerebellum of the dog closely approximates that of the coyote, which is closely aligned with the jackals, and that the wolves show numerous brain traits distinct from the other species (Atkins and Dillon 1971). Wolves also have serological and biochemical traits distinct from dogs (Leone and Wiens 1956; Lauer, Kuyt & Baker 1969).


title: Evolution of the wolf

text: The tails of domestic dogs tend to curl upwards, which is not found in other canid. Dogs possess a domed forehead, which is easy to see in a profile view with a bulge above the orbit (the bony socket) of the eye. The temporalis muscle that closes the jaws is more robust in wolves. Compared with equally sized wolves, dogs possess 20% smaller skulls, and compared with equally weighted wolves dogs possess 20% smaller brains. The tympanic bullae are large, convex, and almost spherical in wolves, while the bullae of dogs are smaller, compressed, and slightly crumpled. Dogs have a distinctive "stop" between the forehead and nose. Domestic dogs are distinguishable from wolves by starch gel electrophoresis of red blood cell acid phosphatase.


title: Evolution of the wolf

text: During the Last Glacial Maximum, there was greater wolf genetic diversity than there is today, and within the Pleistocene grey wolf population the variations between local environments would have encouraged a range of wolf ecotypes that were genetically, morphologically and ecologically distinct from one another. One author has proposed that the most likely explanation for the different morphological characteristics of the dog compared to the wolf is that the dog's ancestor was adapted to a different niche than the wolf.


title: Evolution of the wolf

text: The mitochondria within each cell contain many copies of a small circular DNA genome and in mammals it is 16,000–18,000 base pairs in length. A cell contains hundreds or thousands of mitochondria and therefore the genes contained within those mitochondria are more abundant than the genes that occur in the nucleus of the cell. The abundance of mitochondrial DNA (mDNA) is useful for the genetic analysis of ancient remains where the DNA has degraded.


title: Evolution of the wolf

text: Mitochondrial DNA sequences have a higher mutation rate than the mutation rate of nuclear genes and for mammals this rate is 5–10 times faster. The mitochondrial protein-coding genes evolve much faster and are powerful markers for inferring evolution history at category levels such as families, genera, and species. However, they have evolved at a faster rate than other DNA markers and there is a timing difference in its molecular clock that needs to be validated against other sources. The taxonomic status of uncertain species is better resolved through using nuclear DNA from the nucleus of the cell, which is more suitable for analyzing the recent history. In most cases, mDNA is inherited from the maternal ancestor. Therefore, phylogenetic analysis of mDNA sequences within species provides a history of maternal lineages that can be represented as a phylogenetic tree.


title: Evolution of the wolf

text: The mDNA sequences of the dog and wolf differ by only 0–12 substitutions within 261 base-pairs, whereas dogs always differed from coyotes and jackals by at least 20 substitutions. This finding implies that the dog derived from the wolf and that there has been repeated back-crossing, or that the dog may have descended from a now extinct species of canid whose closest living relative is the modern wolf.


title: Evolution of the wolf

text: Different DNA studies may give conflicting results because of the specimens selected, the technology used, and the assumptions made by the researchers. Any one from a panel of genetic markers can be chosen for use in a study. The techniques used to extract, locate and compare genetic sequences can be applied using advances in technology, which allows researchers to observe longer lengths of base pairs that provide more data to give better phylogenetic resolution. Phylogenetic trees compiled using different genetic markers have given conflicting results on the relationship between the wolf, dog and coyote. One study based on SNPs (a single mutation), and another based on nuclear gene sequences (taken from the cell nucleus), showed dogs clustering with coyotes and separate from wolves. Another study based on SNPS showed wolves clustering with coyotes and separate from dogs. Other studies based on a number of markers show the more widely accepted result of wolves clustering with dogs separate from coyotes. These results demonstrate that caution is needed when interpreting the results provided by genetic markers.


title: Evolution of the wolf

text: There are two key assumptions that are made for dating the divergence time for species: the generation time and the genetic mutation rate per generation. The time between generations for wolves is assumed to be three years based on the extant grey wolf, and two years for the dog based on the extant dog. One recent major study assumed a generation time of 2 years for the dog for as far back as 10,000 years ago, and then assumed a generation time of 3 years (the same as the wolf) before that to calculate a proposed divergence time between the two. In 2017, the wolf research scientist L. David Mech queried why evolutionary biologists were calculating the approximate time of the dog diverging from the wolf through using a wolf generation time of three years when published works using large data sets demonstrate a figure of 4.2–4.7 years. They were encouraged to recalculate their divergence dates accordingly.


title: Evolution of the wolf

text: DNA studies are conducted but with "the mutation rate as the dominant source of uncertainty." In 2005, Lindblad-Toh sequenced the first draft genome of the extant dog, and calculated a proposed mutation rate of 1x10 mutations per generation. In 2015, Skoglund was able to sequence the first draft genome of the 35,000 YBP Taimyr wolf and used its radio-carbon date to validate a proposed genetic mutation rate of 0.4x10 mutations per generation. The difference is a timing factor of 2.5, however another study stated that because only one Pleistocene wolf specimen has so far been sequenced, then the result should be treated with caution, with that study then providing both estimates to calculate the proposed divergence times between the wolf and dog. However, in 2016 the mutation rate of the 4,800 YBP Newgrange dog matched that of the Taimyr wolf.


title: Evolution of the wolf

text: The wolf-like canids (the canid subfamily Caninae) are a group of large carnivores that are genetically closely related because their chromosomes number 78. The group includes genus "Canis", "Cuon" and "Lycaon". The members are the dog "(C. familiaris)", grey wolf ("C. lupus"), coyote ("C. latrans"), golden jackal ("C. aureus"), Ethiopian wolf ("C. simensis"), black-backed jackal ("Lupulella mesomelas"), side-striped jackal ("Lupulella adusta"), dhole ("Cuon alpinus"), and African wild dog ("Lycaon pictus"). Newly proposed members include the red wolf ("Canis rufus"), eastern wolf ("Canis lycaon"), and African golden wolf ("C. anthus"). As they possess 78 chromosomes, all members of the genus "Canis" (coyotes, wolves, jackals) are karyologically indistinguishable from each other, and from the dhole and the African hunting dog. The members of "Canis" can potentially interbreed and there is evidence that the Ethiopian wolf has hybridized with dogs. According to zoologist Reginald Pocock, a dhole interbred with a golden jackal. The African hunting dog is large, highly mobile, known to disperse over large distances and are rare throughout much of their geographical range, making opportunities for hybridization difficult. A study of the maternal mitochondrial DNA of the black-backed jackal could find no evidence of genotypes from the most likely mates – the side-striped jackal nor the golden jackal – indicating that male black-backed jackals had not bred with these. A search of the scientific literature could not find evidence of hybridization for the rare side-striped jackal.


title: Evolution of the wolf

text: A DNA sequence alignment for the wolf-like canids gave a phylogenetic tree with the grey wolf and dog being the most closely related, followed by a close affiliation with the coyote, golden jackal and Ethiopian wolf, and the dog can hybridize in the wild with these three species. Next closest to this group are the dhole and African wild dog that both have unique meat-slicing teeth, suggesting that this adaptation was later lost by the other members. The two African jackals are shown as the most basal members of this clade, which means that this tree is indicating an African origin for the clade. The tree illustrates the genotype–phenotype distinction, where a genotype is an organism's full hereditary information and a phenotype is an organism's actual observed properties, such as morphology, development, or behavior. By phenotype, the dhole (genus "Cuon") and the African hunting dog (genus "Lycaon") are not classified as members of the genus "Canis", but by genotype they are closer to dogs, wolves and coyotes than are the two genus "Canis" jackals – the Side-striped jackal ("C. adustus") and the Black-backed jackal ("C. mesomelas").


title: Evolution of the wolf

text: In 2015, a study of mitochondrial genome sequences and nuclear genome sequences of African and Eurasian canids indicated that extant wolf-like canids had colonized Africa from Eurasia at least 5 times throughout the Pliocene and Pleistocene, which is consistent with fossil evidence suggesting that much of the African canid diversity resulted from the immigration of Eurasian ancestors, likely coincident with Plio-Pleistocene climatic oscillations between arid and humid conditions.


title: Evolution of the wolf

text: The phylogenetic tree for the wolf-like canids may give conflicting positions for the black-backed jackal and the side-striped jackal relative to the genus "Canis" members depending on whether the genetic markers were based on mitochondrial DNA or nuclear DNA. The explanation proposed for this mito-nuclear discord is that mitochondrial DNA introgression occurred from an ancient ancestor of genus "Canis" into the lineage that led to the black-backed jackal around 6.2–5.2 million years ago.


title: Evolution of the wolf

text: Phylogenetic tree of the extant wolf-like canids, with the pink shading representing the species "Canis lupus".


title: Evolution of the wolf

text: In 2018, whole genome sequencing was used to compare members of genus "Canis", along with the dhole ("Cuon alpinus") and the African hunting dog ("Lycaon pictus"). There is evidence of gene flow between African golden wolves, golden jackals, and grey wolves. The study suggests that the African golden wolf is a descendant of a genetically admixed canid of 72% grey wolf and 28% Ethiopian wolf ancestry, and that the Ethiopian wolf once had a wider range in Africa. One African golden wolf from the Egyptian Sinai Peninsula showed high admixture with the Middle Eastern grey wolves and dogs, highlighting the role of the land bridge between the African and Eurasian continents in canid evolution. There is evidence of gene flow between golden jackals and Middle Eastern wolves, less so with European and Asian wolves, and least with North American wolves. The study proposes that the golden jackal ancestry found in North American wolves may have occurred before the divergence of the Eurasian and North American grey wolves. The study indicates that the common ancestor of the coyote and grey wolf has genetically admixed with a ghost population of an extinct unidentified canid. The canid is genetically close to the dhole and has evolved after the divergence of the African hunting dog from the other canid species. The basal position of the coyote compared to the wolf is proposed to be due to the coyote retaining more of the mitochondrial genome of this unknown canid.


title: Evolution of the wolf

text: A genomic study on the wolves of China included museum specimens of wolves from southern China that were collected between 1963 and 1988. The wolves in the study formed 3 clades: north Asian wolves that included those from northern China and eastern Russia, Himalayan wolves from the Tibetan Plateau, and a unique population from southern China. One specimen located as far southeast as Jiangxi province shows evidence of being admixed between Tibetan-related wolves and other wolves in China. One specimen from Zhejiang province in eastern China shared gene flow with the wolves from southern China, however its genome was 12-14 percent admixed with a canid that may be the dhole or an unknown canid that predates the genetic divergence of the dhole. The wolf population from southern China is believed to be still existing in that region.


title: Evolution of the wolf

text: Grey wolves suffered a species-wide population bottleneck (reduction) approximately 25,000 YBP during the Last Glacial Maximum. This was followed by a single population of modern wolves expanding out of a Beringia refuge to repopulate the wolf's former range, replacing the remaining Late Pleistocene wolf populations across Eurasia and North America as they did so. This source population probably did not give rise to dogs, but admixed with dogs which allowed them to gain coat colour genes that are also related to immunity, and provided dogs with genes which allowed them to adapt to high-altitude environments (e.g. Tibet). This suggests that the genetic divergence of European and East Asian dogs could be based on admixture with different sub-populations of wolves.


title: Evolution of the wolf

text: There is little genetic information available on the ancient wolves that existed prior to the bottleneck. However, studies show that one or more of these ancient populations is more directly ancestral to dogs than are modern wolves, and conceivably these were more prone to domestication by the first humans to invade Eurasia.


title: Evolution of the wolf

text: As of 2020, the oldest known intact wolf remains belongs to a mummified pup dated 56,000 YBP that was recovered from the permafrost along a small tributary of Last Chance Creek near Dawson City, Yukon, Canada. A DNA analysis showed that it belonged to the Beringian wolf clade, that the most recent common ancestor of this clade dates to 86,700–67,500 YBP, and that this clade was basal to all other wolves except for the Himalayan wolf.


title: Evolution of the wolf

text: In 2016, a study built on the work of another major study and analyzed the sequences of 12 genes that are located on the heavy strand of the mitochondrial genome of extinct and modern "C. lupus". The study excluded the sequences of the divergent Himalayan wolf and the Indian grey wolf. The ancient specimens were radiocarbon dated and stratagraphically dated, and together with the sequences generated a time-based phylogenetic tree. From the tree, the study was able to infer the most recent common ancestor for all other "C. lupus" specimens – modern and extinct – was 80,000 YBP and this date concurred with the earlier study. The study could find no evidence of a population bottleneck for wolves until a few thousand years ago.


title: Evolution of the wolf

text: The phylogenetic tree showed the polyphyly of American wolves, the Mexican wolf was divergent from other North American wolves, and these other North American wolves formed two closely related clades. A scenario consistent with the phylogenetic, ice sheet and sea-level data was that during the Ice Age when sea levels were at their lowest, there was a single wave of wolf colonization into North America starting with the opening of the Bering land bridge 70,000 YBP and closing during the Late Glacial Maximum of the Yukon corridor that ran through the division between the Laurentide Ice Sheet and the Cordilleran Ice Sheet 23,000 YBP. Mexican wolves were part of the single wave and either diverged from the other wolves before entering North America or once in North America due to the change in its environment.


title: Evolution of the wolf

text: As wolves had been in the fossil record of North America but modern wolves could trace their ancestry back only 80,000 years, the wolf haplotypes that were already in North America were replaced by these invaders, either through competitive displacement or through admixture. The replacement in North America of a basal population of wolves by a more recent one supported the findings of earlier studies. There possibly existed a panmictic wolf population with gene flow spanning Eurasia and North America until the closing of the ice sheets. Once the sheets closed, the southern wolves were isolated and north of the sheets only the Beringian wolf existed. The land bridge became inundated by the sea 10,000 YBP, the sheets receded 12,000–6,000 YBP, the Beringian wolf went extinct and the southern wolves expanded to recolonize the rest of North America. All North American wolves are descended from those that were once isolated south of the ice sheets. However, much of their diversity was later lost during the twentieth century.


title: Evolution of the wolf

text: Studies using mitochondrial DNA have indicated that the wolves of coastal south-east Alaska are genetically distinct from inland grey wolves, reflecting a pattern also observed in other taxa. They show a phylogenetic relationship with extirpated wolves from the south (Oklahoma), indicating that these wolves are the last remains of a once widespread group that has been largely extirpated during the last century, and that the wolves of northern North America had originally expanded from southern refuges below the Wisconsin glaciation after the ice had melted at the end of the Last Glacial Maximum. A whole-genome DNA study indicated that all North American wolves were monophyletic and therefore are the descendants of a common ancestor.


title: Evolution of the wolf

text: During the same period, the Soya Strait between Hokkaido and Sakhalin Island was dry for 75,000 years and it was proposed that the extinct Ezo wolf ("C. l. hattai") arrived on Hokkaido from Sakhalin. However, the sequences indicated that it arrived in Hokkaido less than 10,000 YBP. The Ezo wolf was closely related to one of the North American clades, but different from the more southerly Japanese wolf ("C. l. hodophilax") that was basal to modern wolves. The Japanese wolf inhabited Kyushu, Shikoku, and Honshu islands but not Hokkaido Island. This indicates that its ancestor may have migrated from the Asian continent through the Korean Peninsula into Japan. The past sea levels of the Korean Strait together with the timing of the Japanese wolf sequences indicated that it arrived to the southern islands less than 20,000 YBP. In 2020, a genomic study found that the Japanese wolf was the last of the Siberian Pleistocene wolves, which were thought to have gone extinct at the end of the Late Pleistocene (11,700 years ago). Some of these had survived into the 20th Century and had admixed with Japanese dogs.


title: Evolution of the wolf

text: The dog was a very successful invader of North America and had established a widespread ecological niche by the Early–Middle Holocene. There was no overlap in niche between the dog and the wolf in comparison to the dog and other North American canids. By the Late Holocene, the dog's niche area was less in size than researchers had expected to find, indicating that it was limited by biotic factors. These regions include the northeast and northwest of the United States that correlate with the greatest densities of early human occupation, indicating that the dog had "defected" from the wolf niche to the human niche and explains why the dog's niche area was not as large as expected. The separation between dog and wolf may reflect the rapid rate in which domestication occurred, including the possibility of a second domestication event occurring in North America. Packs of wolves and hunter-gatherers hunt similar prey in a similar way within a similar group social structure that may have facilitated wolf domestication.


title: Evolution of the wolf

text: The wolf was exterminated in the southern part of their historic geographical range in North America by the middle of the 20th century. An mDNA study of 34 wolf remains from North America dated between 1856 and 1915 found their genetic diversity to be twice that of modern wolves in these regions, and two thirds of the haplotypes identified were unique. These results indicate that a historic population of several hundred thousand wolves once existed in Mexico and the western US.


title: Evolution of the wolf

text: In 1993, a study proposed that the wolves of North America display skull traits more similar towards the coyote than those wolves from Eurasia. In 2016, a whole-genome DNA study proposed, based on the assumptions made, that all of the North American wolves and coyotes diverged from a common ancestor less than 6,000–117,000 years ago. The study also indicated that all North American wolves have a significant amount of coyote ancestry and all coyotes some degree of wolf ancestry, and that the red wolf and eastern wolf are highly admixed with different proportions of grey wolf and coyote ancestry. However, the test that was used cannot differentiate between anceint and recent hybridization. One test indicated a wolf/coyote divergence time of 51,000 years before present that matched other studies indicating that the extant wolf came into being around this time. Another test indicated that the red wolf diverged from the coyote between 55,000–117,000 years before present and the Great Lakes region wolf 32,000 years before present. But a 2000 study on the haplotypes of North American canids showed that the divergences between coyotes, grey wolves, red wolves, and algonquin (eastern) wolves happened much earlier. This study showed that there was a 3.2 % divergence between the eastern Canadian wolf and coyote gene sequences, and a 2.3 % sequence divergence between the red wolf and coyote haplotypes. However, the divergence was a much greater 8.0% between gray wolf (C. lupus) mtDNA and eastern/red wolf haplotypes, and 10.0% between gray wolf and coyote haplotypes. The sequence difference observed between eastern Canadian wolf sequences and coyote sequences is consistent with a separation of 150 000 – 300 000 years, using a divergence rate of 1–2% per 100 000 years. Additionally, conflicting with the recent divergent hypothesis, the fossil record that indicates a coyote-like specimen dated to 1 million years before present.


title: Evolution of the wolf

text: The modern grey wolf expanded out of Beringia 25,000 years ago. The modern coyote appeared around 10,000 years ago, and most likely descended from the Pleistocene coyote. The most genetically basal coyote mDNA clade pre-dates the Last Glacial Maximum and is a haplotype that can only be found in the Eastern wolf. This implies that the large, wolf-like Pleistocene coyote was the ancestor of the Eastern wolf and Red wolf, which diverged from each other later in time.


title: Evolution of the wolf

text: The domestic dog ("Canis lupus familiaris") is the most widely abundant large carnivore. Over the past million years, numerous wolf-like forms existed but their turnover has been high, and modern wolves are not the lineal ancestors of dogs. Although research had suggested that dogs and wolves were genetically very close relatives, later phylogenetic analysis strongly supported the hypothesis that dogs and wolves are reciprocally monophylic taxa that form two sister clades. This suggests that none of the modern wolf populations are related to the wolves that were first domesticated and the wolf ancestor of dogs is therefore presumed extinct. Recent mitochondrial DNA analyses of ancient and modern grey wolf specimens supports a pattern of population reduction and turnover. An alternate proposal is that during the ecological upheavals of the Late Pleistocene all of the remaining members of a dwindling lineage joined humans.