The world's oldest captive white rhino, South African-born Sana, has died at the age of 55, the French zoo that she called home for the last 26 years said Thursday.
Born in 1964 in the Umfolozi National Park in South Africa's eastern KwaZulu-Natal province, the southern white rhino was transported to Europe seven years later as a zoo attraction.
Sana lived in several parks in Germany before arriving in 1993 at Planete Sauvage, a zoo in western France near the city of Nantes that had opened the previous year.
The life expectancy of white rhinos in the wild is about 50 years, according to the Save the Rhino advocacy group.
Sana required special care in her later years, the park said in a statement.
Notably, she was no longer able to roll in the mud, a popular and crucial pastime that helps rhinos regulate their body temperature, prevent sunburn and ward off pesky insects.
"Once a week, they coated her in green clay to help her skin stay hydrated and to stave off infections," the park said.
It added that Sana "always had a strong character, she was the one who set the rules."
Her death has "deeply affected" her caretakers, the zoo said.
Adult white rhinos weigh between 1.8 and 2.5 tonnes (5,600 pounds), according to Save the Rhino.
Their natural habitats are tropical and subtropical grasslands, savannas and shrublands found in South Africa, Botswana, Kenya, Namibia, Swaziland, Zambia, Zimbabwe and Uganda.
The square-lipped grazers, larger than their black cousins, are divided into southern and northern subspecies.
The last male northern white rhino, Sudan, died in Kenya last year, leaving two sole captive female survivors, neither able to carry a calf.
No northern white rhinos are left in the wild.
In September, conservationists created two northern white rhino embryos from the two last females and sperm from deceased males, to one day implant into a surrogate mother in the hope of reviving the line.
Save the Rhino says around 20,000 southern white rhinos -- Sana's tribe -- are left in the world.
Black rhinos, however, are considered critically endangered, with only about 5,000 left.
Rhino horn is a much sought-after ingredient in Asian traditional medicine.
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