The grandmother of Harambe, the gorilla infamously shot dead by zookeepers after a toddler wandered into his enclosure on 28 May 2016, was put down in Miami on 18 December. Josephine, a 49-year-old matriarch, was suffering from complex health problems and was struggling to move – leaving staff no option but to "humanely euthanise" her.

Her grandson Harambe had just turned 17 when he was gunned down at Cincinnati Zoo. The adolescent gorilla was carrying a small boy that had crawled into his enclosure.

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The Western lowland gorilla's death sparked global controversy with many criticising the zoo's decision to kill him because, they claim, he was not behaving aggressively. Others argued that the incident served to highlight the ethical problems posed by the very existence of zoos.

The boy's mother, Michelle Gregg, received a tirade of abuse for allowing her child to climb a three-foot-high wall, crawl through a bush and fall 15ft into the gorilla enclosure. Some animal-rights activists blamed her for Harambe's death.

Gregg responded to the criticism, saying: "As a society, we are quick to judge how a parent could take their eyes off of their child, and if anyone knows me I keep a tight watch on my kids. Accidents happen, but I am thankful that the right people were in the right place today."

Harambe's grandmother Josephine was born arrived at Miami Zoo in 1983. In 1984 she gave birth to Moja, who would father Harambe in Texas. The death of Josephine comes just a week after Binta, the first chimpanzee to be born at the zoo, died of a blood clot, according to NBC Miami.