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Dian Fossey (; January 16, 1932 - c. December 26, 1985) was an American zoologist, primatologist, and anthropologist who undertook an extensive study of mountain gorilla groups over a period of 18 years. She studied them daily in the mountain forests of Rwanda, initially encouraged to work there by anthropologist Louis Leakey. Her 1983 book, Gorillas in the Mist, combines her scientific study of the gorillas at Karisoke Research Center with her own personal story. It was adapted into a 1988 film of the same name.

Called one of the foremost primatologists in the world, Fossey, along with Jane Goodall and Birut? Galdikas, were the so-called Trimates, a group of three prominent researchers on primates (Fossey on gorillas; Goodall on common chimpanzees; and Galdikas on orangutans) sent by Leakey to study great apes in their natural environments.


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Life and career

Fossey was born in San Francisco, California, the daughter of Kathryn "Kitty" (née Kidd), a fashion model, and George E. Fossey III, an insurance agent. Her parents divorced when she was six. Her mother remarried the following year, to businessman Richard Price. Her father tried to keep in full contact, but her mother discouraged it, and all contact was subsequently lost. Fossey's stepfather, Richard Price, never treated her as his own child. He would not allow Fossey to sit at the dining room table with him or her mother during dinner meals. A man adhering to strict discipline, Richard Price offered Fossey little to no emotional support. Struggling with personal insecurity, Fossey turned to animals as a way to gain acceptance. Her love for animals began with her first pet goldfish and continued throughout her entire life. At age six, she began horse riding, earning a letter from her school; by her graduation in 1954, Fossey had established herself as an equestrienne.

Education

Educated at Lowell High School, following the guidance of her stepfather she enrolled in a business course at the College of Marin. However, spending her summer on a ranch in Montana at age 19 rekindled her love of animals, and she enrolled in a pre-veterinary course in biology at the University of California, Davis. In defiance to her stepfather's wishes that she attend a business school, Dian wanted to spend her professional life working with animals. As a consequence, Dian's parents failed to give her any substantial amount of financial support throughout her adult life. She supported herself by working as a clerk at White Front (a department store), doing other clerking and laboratory work, and laboring as a machinist in a factory.

Although Fossey had always been an exemplary student, she had difficulties with basic sciences including chemistry and physics, and failed her second year of the program. She transferred to San Jose State College where she was a member of Kappa Alpha Theta sorority to study occupational therapy, receiving her bachelor's degree in 1954. Initially following her college major, Fossey began a career in occupational therapy. She interned at various hospitals in California and worked with tuberculosis patients. Fossey was originally a prizewinning equestrian, which drew her to Kentucky in 1955, and a year later took a job as an occupational therapist at the Kosair Crippled Children's Hospital in Louisville.

Her shy and reserved personality allowed her to work well with the children at the hospital. Fossey became close with her coworker Mary White "Gaynee" Henry, secretary to the hospital's chief administrator and the wife of one of the doctors, Michael J. Henry. The Henrys invited Fossey to join them on their family farm, where she worked with livestock on a daily basis and also experienced an inclusive family atmosphere that had been missing for most of her life. During her free time she would pursue her love of horses.

Interest in Africa

Fossey turned down an offer to join the Henrys on an African tour due to lack of finances, but in 1963 she borrowed $8,000 (one year's salary), took out her life savings and went on a seven-week visit to Africa. In September 1963, she arrived in Nairobi, Kenya. While there, she met actor William Holden, owner of Treetops Hotel, who introduced her to her safari guide, John Alexander. Alexander became her guide for the next seven weeks through Kenya, Tanzania, Democratic Republic of Congo, and Rhodesia. Alexander's route included visits to Tsavo, Africa's largest national park; the saline lake of Manyara, famous for attracting giant flocks of flamingos; and the Ngorongoro Crater, well known for its abundant wildlife. The final two sites for her visit were Olduvai Gorge in Tanzania (the archeological site of Louis and Mary Leakey); and Mt. Mikeno in Congo, where in 1959, American zoologist George Schaller had carried out a yearlong pioneering study of the mountain gorilla. At Olduvai Gorge, Fossey met the Leakeys while they were examining the area for hominid fossils. Leakey talked to Fossey about the work of Jane Goodall and the importance of long-term research of the great apes. Although she had broken her ankle while visiting the Leakeys, by October 16, Fossey was staying in Walter Baumgartel's small hotel in Uganda, the Travellers Rest. Baumgartel, an advocate of gorilla conservation, was among the first to see the benefits that tourism could bring to the area, and he introduced Fossey to Kenyan wildlife photographers Joan and Alan Root. The couple agreed to allow Fossey and Alexander to camp behind their own camp, and it was during these few days that Fossey first encountered wild mountain gorillas. After staying with friends in Rhodesia, Fossey returned home to Louisville to repay her loans. She published three articles in The Courier-Journal newspaper, detailing her visit to Africa.

Research in the Congo

When Leakey made an appearance in Louisville while on a nationwide lecture tour, Fossey took the color supplements that had appeared about her African trip in The Courier-Journal to show to Leakey, who remembered her and her interest in mountain gorillas. Three years after the original safari, Leakey suggested that Fossey could undertake a long-term study of the gorillas in the same manner as Jane Goodall had with chimpanzees in Tanzania. Leakey lined up funding for Fossey to research mountain gorillas, and Fossey left her job to relocate to Africa.

After studying Swahili and auditing a class on primatology (the scientific study of primates) during the eight months it took to get her visa and funding, Fossey arrived in Nairobi in December 1966. With the help of Joan Root and Leakey, Fossey acquired the necessary provisions and an old canvas-topped Land Rover which she named "Lily". On the way to the Congo, Fossey visited the Gombe Stream Research Centre to meet Goodall and observe her research methods with chimpanzees. Accompanied by photographer Alan Root, who helped her obtain work permits for the Virunga Mountains, Fossey began her field study at Kabara, in the Congo in early 1967, in the same meadow where Schaller had made his camp seven years earlier. Root taught her basic gorilla tracking, and his tracker Sanwekwe later helped in Fossey's camp. Living in tents on mainly tinned produce, once a month Fossey would hike down the mountain to "Lily" and make the two-hour drive to the village of Kikumba to restock.

Fossey identified three distinct groups in her study area, but could not get close to them. She eventually found that mimicking their actions and making grunting sounds assured them, together with submissive behavior and eating of the local celery plant. She later attributed her success with habituating gorillas to her experience working as an occupational therapist with autistic children. Like George Schaller, Fossey relied greatly on individual "noseprints" for identification, initially via sketching and later by camera.

Fossey had arrived in the Congo in locally turbulent times. Known as the Belgian Congo until its independence in June 1960, unrest and rebellion plagued the new government until 1965, when Lieutenant General Joseph-Désiré Mobutu, by then commander-in-chief of the national army, seized control of the country and declared himself president for five years during what is now called the Congo Crisis. During the political upheaval, a rebellion and battles took place in the Kivu Province. On July 9, 1967, soldiers arrived at the camp to escort Fossey and her research workers down, and she was interned at Rumangabo for two weeks. Fossey eventually escaped through bribery to Walter Baumgärtel's Travellers Rest Hotel in Kisoro, where her escort was arrested by the Ugandan military. Advised by the Ugandan authorities not to return to Congo, after meeting Leakey in Nairobi, Fossey agreed with him against US Embassy advice to restart her study on the Rwandan side of the Virungas. In Rwanda, Fossey had met local American expatriate Rosamond Carr, who introduced her to Belgian local Alyette DeMunck; DeMunck had a local's knowledge of Rwanda and offered to find Fossey a suitable site for study.


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Conservation work in Rwanda

On September 24, 1967, Fossey founded the Karisoke Research Center, a remote rainforest camp nestled in Ruhengeri province in the saddle of two volcanoes. For the research center's name, Fossey used "Kari" for the first four letters of Mount Karisimbi that overlooked her camp from the south, and "soke" for the last four letters of Mount Bisoke, the slopes of which rose to the north, directly behind camp. Established 3,000 metres (9,800 ft) up Mount Bisoke, the defined study area covered 25 square kilometres (9.7 sq mi). She became known by locals as Nyirmachabelli, or Nyiramacibiri, roughly translated as "The woman who lives alone on the mountain."

Unlike the gorillas from the Congo side of the Virungas, the Karisoke area gorillas had never been partially habituated by Schaller's study; they knew humans only as poachers, and it took longer for Fossey to be able to study the Karisoke gorillas at a close distance.

Many research students left after not being able to handle the cold, dark, and extremely muddy conditions around Karisoke on the slopes of the Virunga Volcanoes, where paths usually had to be cut through six-foot-tall grass with a machete.

Opposition to poaching

While hunting had been illegal in the national park of the Virunga Volcanoes in Rwanda since the 1920s, the law was rarely enforced by park conservators, who were often bribed by poachers and paid a salary less than Fossey's own African staff. On three occasions, Fossey wrote that she witnessed the aftermath of the capture of infant gorillas at the behest of the park conservators for zoos; since gorillas will fight to the death to protect their young, the kidnappings would often result in up to 10 adult gorillas' deaths. Through the Digit Fund, Fossey financed patrols to destroy poachers' traps in the Karisoke study area. In four months in 1979, the Fossey patrol consisting of four African staffers destroyed 987 poachers' traps in the research area's vicinity. The official Rwandan national park guards, consisting of 24 staffers, did not eradicate any poachers' traps during the same period. In the eastern portion of the park not patrolled by Fossey, poachers virtually eradicated all the park's elephants for ivory and killed more than a dozen gorillas.

Fossey helped in the arrest of several poachers, some of whom served or are serving long prison sentences.

In 1978, Fossey attempted to prevent the export of two young gorillas, Coco and Pucker, from Rwanda to the zoo in Cologne, Germany. During the capture of the infants at the behest of the Cologne Zoo and Rwandan park conservator, 20 adult gorillas had been killed. The infant gorillas were given to Fossey by the park conservator of the Virunga Volcanoes for treatment of injuries suffered during their capture and captivity. With considerable effort, she restored them to some approximation of health. Over Fossey's objections, the gorillas were shipped to Cologne, where they lived nine years in captivity, both dying in the same month. She viewed the holding of animals in "prison" (zoos) for the entertainment of people as unethical.

While gorillas from rival gang groups on the mountains that were not part of Fossey's study had often been found poached five to ten at a time, and had spurred Fossey to conduct her own anti-poaching patrols, Fossey's study groups had not been direct victims of poaching until Fossey's favorite gorilla Digit was killed in 1978. Later that year, the silverback of Digit's Group 4, named for Fossey's Uncle Bert, was shot in the heart while trying to save his son, Kweli, from being seized by poachers cooperating with the Rwandan park conservator. Kweli's mother, Macho, was also killed in the raid, but Kweli was not captured due to Uncle Bert's intervention; however, three-year-old Kweli died slowly and painfully of gangrene, from being brushed by a poacher's bullet.

According to Fossey's letters, ORTPN (the Rwandan national park system), the World Wildlife Fund, African Wildlife Foundation, Fauna Preservation Society, the Mountain Gorilla Project and some of her former students tried to wrest control of the Karisoke research center from her for the purpose of tourism, by portraying her as unstable. In her last two years, Fossey claims not to have lost any gorillas to poachers; however, the Mountain Gorilla Project, which was supposed to patrol the Mount Sabyinyo area, tried to cover up gorilla deaths caused by poaching and diseases transmitted through tourists. Nevertheless, these organizations received most of the public donations directed toward gorilla conservation. The public often believed their money would go to Fossey, who was struggling to finance her anti-poaching and bushmeat hunting patrols, while organizations collecting in her name put it into tourism projects and as she put it "to pay the airfare of so-called conservationists who will never go on anti-poaching patrols in their life." Fossey described the differing two philosophies as her own "active conservation" or the international conservation groups' "theoretical conservation."

Opposition to tourism

Fossey strongly opposed wildlife tourism, as gorillas are very susceptible to human anthroponotic diseases like influenza for which they have no immunity. Fossey reported several cases in which gorillas died because of diseases spread by tourists. She also viewed tourism as an interference into their natural wild behavior. Fossey also criticized tourist programs, often paid for by international conservation organizations, for interfering with both her research and the peace of the mountain gorillas' habitat, and was concerned Jane Goodall, who actually joined a chimpanzee society as a member, was inappropriately changing her study subjects' behavior.

Today, however, the Dian Fossey Gorilla Fund International promotes tourism, which they say helps to create a stable and sustainable local community dedicated to protecting the gorillas and their habitat.

Preservation of habitat

Fossey is responsible for the revision of a European Community project that converted parkland into pyrethrum farms. Thanks to her efforts, the park boundary was lowered from the 3,000-meter line to the 2,500-meter line.

Digit Fund

Sometime during the day on New Year's Eve 1977, Fossey's favorite gorilla, Digit, was killed by poachers. As the sentry of study group 4, he defended the group against six poachers and their dogs, who ran across the gorilla study group while checking antelope traplines. Digit took five spear wounds in ferocious self-defence and managed to kill one of the poachers' dogs, allowing the other 13 members of his group to escape. Poachers sell gorilla hands as delicacies, magic charms or to make ash trays. Digit was decapitated, and his hands cut off for ashtrays, for the price of $20. After his mutilated body was discovered by research assistant Ian Redmond, Fossey's group captured one of the killers. He revealed the names of his five accomplices, three of whom were later imprisoned.

Fossey subsequently created the Digit Fund (now the Dian Fossey Gorilla Fund International in the USA) to raise money for anti-poaching patrols. In addition, a consortium of international gorilla funds arose to accept donations in light of Digit's death and increased attention on poaching. Fossey mostly opposed the efforts of the international organizations, which she felt inefficiently directed their funds towards more equipment for Rwandan park officials, some of whom were alleged to have ordered some of the gorilla poachings in the first place.

The deaths of some of her most studied gorillas caused Fossey to devote more of her attention to preventing poaching and less on scientific publishing and research. Fossey became more intense in protecting the gorillas and began to employ more direct tactics: she and her staff cut animal traps almost as soon as they were set; frightened, captured and humiliated the poachers; held their cattle for ransom; burned their hunting camps and even mats from their houses. Fossey also constantly challenged the local officials to enforce the law and assist her.


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Personal life

During her African safari, Fossey met Alexie Forrester, the brother of an African she had been dating in Louisville; Fossey and Forrester later became engaged. In her later years, Fossey became involved with National Geographic photographer Bob Campbell after a year of working together at Karisoke, with Campbell promising to leave his wife. Eventually the pair grew apart through her dedication to the gorillas and Karisoke, along with his need to work further afield and his marriage. In 1970, during her time at Darwin College, Cambridge under the University of Cambridge to get her Ph.D., she discovered she was pregnant and had an abortion, later commenting that "you can't be a cover girl for National Geographic magazine and be pregnant." Nevertheless, she graduated with a Doctor of Philosophy in Zoology in 1976. Fossey had other relationships throughout the years and always had a love for children. Since Fossey would rescue any abused or abandoned animal she saw in Africa or near Karisoke, she acquired a menagerie in the camp, including a monkey who lived in her cabin, Kima, and a dog, Cindy. Fossey held Christmas parties every year for her researchers, staffers, and their families, and she developed a genuine friendship with Jane Goodall.

Fossey had been plagued by lung problems from an early age, and later in her life, she suffered from advanced emphysema brought on by years of heavy cigarette smoking. As the debilitating disease progressed--further aggravated by the high mountain altitude and damp climate--Fossey found it increasingly difficult to conduct field research, frequently suffering from shortness of breath and requiring the help of an oxygen tank when climbing or hiking long distances.


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Death

In the early morning of December 27, 1985, Fossey was discovered murdered in the bedroom of her cabin located at the far edge of the camp in the Virunga Mountains, Rwanda. Her body was found face-up near the two beds where she slept, roughly 7 feet (2 m) away from a hole that her assailant(s) had apparently cut in the wall of the cabin. Wayne Richard McGuire, Fossey's last research assistant at Karisoke, was summoned to the scene by Fossey's house servant and found her bludgeoned to death, reporting that "when I reached down to check her vital signs, I saw her face had been split, diagonally, with one machete blow." The cabin was littered with broken glass and overturned furniture, with a 9-mm handgun and ammunition beside her on the floor. Robbery was not believed to be the motive for the crime, as Fossey's valuables were still in the cabin, including her passport, handguns, and thousands of dollars in U.S. bills and traveler's checks.

The last entry in her diary read:

When you realize the value of all life, you dwell less on what is past and concentrate more on the preservation of the future.

Fossey is buried at Karisoke, in a site that she herself had constructed for her deceased gorilla friends. She was buried in the gorilla graveyard next to Digit, and near many gorillas killed by poachers. Memorial services were also held in New York, Washington, and California.

A will purporting to be Fossey's bequeathed all of her estate (including the proceeds from the film Gorillas in the Mist) to the Digit Fund to underwrite anti-poaching patrols. Fossey did not mention her family in the will, which was unsigned. Her mother, Hazel Fossey Price, challenged the will and was successful. Supreme Court Justice Swartwood threw out the will and awarded the estate to her mother, including about $4.9 million in royalties from a recent book and upcoming movie, stating that the document "was simply a draft of her purported will and not a will at all." Price said she was working on a project to preserve the work their daughter had done for the mountain gorillas in Rwanda, located in eastern central Africa south of Uganda.

Aftermath

After Fossey's death, her entire staff, including Rwelekana, a tracker she had fired months before, were arrested. All were later released except Rwelekana, who was later found dead in prison, supposedly having hanged himself.

Rwandan courts later tried and convicted Wayne McGuire in absentia for her murder. McGuire had returned to the United States in July 1987, and because no extradition treaty exists between the U.S. and Rwanda, McGuire, whose guilt is still widely questioned, has not served his sentence.

Following his return to the U.S., McGuire gave a brief statement at a news conference in Century City, Los Angeles, saying Fossey had been his "friend and mentor", calling her death "tragic" and the charges "outrageous". Thereafter, McGuire was largely under the radar until 2005, when news broke that he had been accepted for a job with the Health and Human Services division of the State of Nebraska. The job offer was revoked upon discovery of his relation to the Fossey case.

Several subsequent books, including Farley Mowat's biography of Fossey, Woman in the Mists (New York, NY: Warner Books, 1987), have suggested alternate theories regarding her murder including intimations that she may have been killed by financial interests linked to tourism or illicit trade.


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Controversy

Fossey was reported to have captured and held Rwandans whom she suspected of poaching, stripped and then beaten them with stinging nettles. After her murder, Fossey's National Geographic editor, Mary Smith, told Shlachter that on visits to the United States, Fossey would "load up on firecrackers, cheap toys and magic tricks as part of her method to mystify the (Africans) -- hold them at bay."

Writing in The Wall Street Journal in 2002, Tunku Varadarajan described Fossey at the end of her life as colorful, controversial, and "a racist alcoholic who regarded her gorillas as better than the African people who lived around them."


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Scientific achievements

Fossey made discoveries about gorillas including how females transfer from group to group over the decades, gorilla vocalization, hierarchies and social relationships among groups, rare infanticide, gorilla diet, and how gorillas recycle nutrients. Fossey's research was funded by the Wilkie Foundation and the Leakey Home, with primary funding from the National Geographic Society.

By 1980, Fossey, who had obtained her PhD at Cambridge University in the UK, was recognized as the world's leading authority on the physiology and behavior of mountain gorillas, defining gorillas as being "dignified, highly social, gentle giants, with individual personalities, and strong family relationships."

Fossey lectured as professor at Cornell University in 1981-83. Her bestselling book Gorillas in the Mist was praised by Nikolaas Tinbergen, the Dutch ethologist and ornithologist who won the 1973 Nobel Prize in Physiology or Medicine. Her book remains the best-selling book about gorillas.


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Legacy

After her death, Fossey's Digit Fund in the U.S. was renamed the Dian Fossey Gorilla Fund International. The Karisoke Research Center is operated by the Dian Fossey Gorilla Fund International, and continues the daily gorilla monitoring and protection that she started.

Shirley McGreal, a friend of Fossey, continues to work for the protection of primates through the work of her International Primate Protection League (IPPL) one of the few wildlife organizations that according to Fossey effectively promotes "active conservation".

Between Fossey's death until the 1994 Rwandan genocide, Karisoke was directed by former students, some of whom had opposed her. During the genocide and subsequent period of insecurity, the camp was completely looted and destroyed. Today only remnants remain of her cabin. During the civil war, the Virunga National Park was filled with refugees, and illegal logging destroyed vast areas.

The Rwandan people adapted the traditional household baby naming ceremony Kwita Izina to use with the gorillas.

Her 82nd birthday in 2014 was marked by a Google Doodle appearing on its search homepage worldwide.

Biographies

Mowat's Virunga, whose British and U.S. editions are called Woman in the Mists: The Story of Dian Fossey and the Mountain Gorillas of Africa, was the first book-length biography of Fossey, and it serves as an insightful counterweight to the many omissions in Fossey's own story, being derived from Fossey's actual letters and entries in her journals. Harold Hayes's book The Dark Romance of Dian Fossey was published in 1989 after extensive interviews with people who lived and worked with Fossey. Haye's book shows Fossey in a less positive or romanticized light than previous accounts had done. The film Gorillas in the Mist was based on Hayes' 1987 article in Life magazine, as cited in the film's credits, instead of Fossey's self-edited autobiography by that title.

No One Loved Gorillas More (2005) was written by Camilla de la Bedoyere and published by National Geographic in the United States and Palazzo Editions in the United Kingdom. Gorilla Dreams: The Legacy of Dian Fossey was written by the investigative journalist Georgianne Nienaber and published in 2006. This account of Fossey's story is told as if in her own words from beyond the grave. Fossey is also prominently featured in a book by Vanity Fair journalist Alex Shoumatoff called African Madness, in which the author expands on Fossey's controversial behaviors, implying that Fossey provoked her own murder by way of her private and public inflammatory interactions with people. The author also wrote a lengthy article titled "The Fatal Obsession of Dian Fossey".

Films, television, and an opera

The Kentucky Opera Visions Program, in Louisville, has written an opera about Fossey, entitled Nyiramachabelli; it premiered on May 23, 2006.

Universal Studios bought the film rights to Gorillas in the Mist from Fossey in 1985, and Warner Bros. Studios bought the rights to the Hayes article, despite its having been severely criticized by Rosamond Carr. As a result of a legal battle between the two studios, a co-production was arranged. Portions of the story and the Hayes article were adapted for the film Gorillas in the Mist, starring Sigourney Weaver. The book covers Fossey's scientific career in great detail and omits material on her personal life, such as her affair with photographer Bob Campbell. In the film, the affair with Campbell (played by Bryan Brown) forms a major subplot. The Hayes article preceding the movie portrayed Fossey as a woman obsessed with gorillas, who would stop at nothing to protect them. The film includes scenes of Fossey's ruthless dealings with poachers, including a scene in which she sets fire to a poacher's home.

In the 2011 BBC documentary All Watched Over by Machines of Loving Grace, Adam Curtis uses Fossey as a symbol of the ideology of ecology, a balance of nature and western post-colonial political exploits in Africa.

"Wild Kingdom: Reunion With the Gorillas" (1984), features footage of Dr. Fossey interacting with the great apes in the Virunga Mountains after being away from them for about three years. The documentary-style video is narrated by Dian Fossey and Marlin Perkins. She shares some of her observations and personal experiences, concerning the gorillas in the band.


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Selected bibliography

Books

  • Dian Fossey: Gorillas in the Mist, Houghton Mifflin Company, 1983
  • "Living with mountain gorillas", in The Marvels of Animal Behavior 208-229 (T.B. Allen ed., National Geographic Society), 1972
  • D. Fossey & A.H. Harcourt: "Feeding ecology of free-ranging mountain gorilla (Gorilla gorilla beringei)", in Primate Ecology: Studies of Feeding and Ranging Behaviour in Lemurs, Monkeys and Apes 415-447 (Tim Clutton-Brock ed., Academic Press), 1977
  • "Development of the mountain gorilla (Gorilla gorilla beringei) through the first thirty-six months", in The Great Apes 139-186 (D.A. Hamburg & E.R. McCown eds., Benjamin-Cummings), 1979

Scholarly articles

  • "An amiable giant: Fuertes's gorilla", Living Bird Quarterly 1(summer): 21-22, 1982.
  • "Mountain gorilla research, 1974", Nat. Geogr. Soc. Res. Reps. 14: 243-258, 1982
  • "Mountain gorilla research, 1971-1972", Nat. Geogr. Soc. Res. Reps. 1971 Projects, 12: 237-255, 1980
  • "Mountain gorilla research, 1969-1970", Nat. Geogr. Soc. Res. Reps. 1969 Projects, 11: 173-176, 1978
  • The behaviour of the mountain gorilla, Ph.D. diss. Cambridge University, 1976
  • "Observations on the home range of one group of mountain gorillas (Gorilla gorilla beringei)", Anim. Behav. 22: 568-581, 1974
  • "Vocalizations of the mountain gorilla (Gorilla gorilla beringei)", Anim. Behav. 20: 36-531972

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