A rodent-borne virus led to the death of the only African elephant at the zoo in Indian capital Delhi, officials have told the BBC.

Shankar, a 29-year-old male elephant, died on 17 September after spending much of his life in isolation. The cause of his death wasn't known at the time.

Now, an autopsy has revealed that the elephant tested positive for encephalomyocarditis virus (EMCV), Delhi Zoo director Sanjeet Kumar said. EMCV is known to cause fatal inflammation of the heart and sometimes brain fever in mammals. The virus is spread through rodent faeces and urine.

Most outbreaks of EMCV have been associated with captive animals in pig farms, primate research centres and zoos. The virus replicates rapidly and can attack the heart and sometimes the brain, often causing sudden death. There is no specific antiviral cure available for EMCV so far.

The virus was first isolated in 1945 from a gibbon in Florida, and since the 1970s, outbreaks have been reported in the US, South Africa, China, Australia, Canada, South America and several countries in Europe. In India, the virus was first isolated in the late 1960s, but Shankar's case is the first recorded death caused by EMCV in the country.

Shankar's death has sparked sadness among animal lovers and activists who had long sought to rehabilitate the lonely animal. He was among two African elephants that arrived in India in 1998 as a diplomatic gift from Zimbabwe. After his companion died in 2001, Shankar was temporarily lodged with Asian elephants, but this arrangement did not work out.

In 2012, he was moved to a new enclosure, essentially confining him to solitude, despite a 2009 federal ban on keeping elephants alone for more than six months. Activists have demanded his relocation to a wildlife sanctuary that houses other African elephants.

After Shankar's death, only one African elephant remains in India, an adult male at Mysore Zoo, who has also been living alone for years.